Automatic tracking of medial gastrocnemius fascicle length during human locomotion

During human locomotion lower extremity muscle-tendon units undergo cyclic length changes that were previously assumed to be representative of muscle fascicle length changes. Measurements in cats and humans have since revealed that muscle fascicle length changes can be uncoupled from those of the muscle-tendon unit. Ultrasonography is frequently used to estimate fascicle length changes during human locomotion. Fascicle length analysis requires time consuming manual methods that are prone to human error and experimenter bias. To bypass these limitations, we have developed an automatic fascicle tracking method based on the Lucas-Kanade optical flow algorithm with an affine optic flow extension. The aims of this study were to compare gastrocnemius fascicle length changes during locomotion using the automated and manual approaches and to determine the repeatability of the automated approach. Ultrasound was used to examine gastrocnemius fascicle lengths in eight participants walking at 4, 5, 6, and 7 km/h and jogging at 7 km/h on a treadmill. Ground reaction forces and three dimensional kinematics were recorded simultaneously. The level of agreement between methods and the repeatability of the automated method were quantified using the coefficient of multiple correlation (CMC). Regardless of speed, the level of agreement between methods was high, with overall CMC values of 0.90 ± 0.09 (95% CI: 0.86-0.95). Repeatability of the algorithm was also high, with an overall CMC of 0.88 ± 0.08 (95% CI: 0.79-0.96). The automated fascicle tracking method presented here is a robust, reliable, and time-efficient alternative to the manual analysis of muscle fascicle length during gait.     

Automated regional analysis of B-mode ultrasound images of skeletal muscle movement

To understand the functional significance of skeletal muscle anatomy, a method of quantifying local shape changes in different tissue structures during dynamic tasks is required. Taking advantage of the good spatial and temporal resolution of B-mode ultrasound imaging, we describe a method of automatically segmenting images into fascicle and aponeurosis regions and tracking movement of features, independently, in localized portions of each tissue. Ultrasound images (25 Hz) of the medial gastrocnemius muscle were collected from eight participants during ankle joint rotation (2° and 20°), isometric contractions (1, 5, and 50 Nm), and deep knee bends. A Kanade-Lucas-Tomasi feature tracker was used to identify and track any distinctive and persistent features within the image sequences. A velocity field representation of local movement was then found and subdivided between fascicle and aponeurosis regions using segmentations from a multiresolution active shape model (ASM). Movement in each region was quantified by interpolating the effect of the fields on a set of probes. ASM segmentation results were compared with hand-labeled data, while aponeurosis and fascicle movement were compared with results from a previously documented cross-correlation approach. ASM provided good image segmentations (<1 mm average error), with fully automatic initialization possible in sequences from seven participants. Feature tracking provided similar length change results to the cross-correlation approach for small movements, while outperforming it in larger movements. The proposed method provides the potential to distinguish between active and passive changes in muscle shape and model strain distributions during different movements/conditions and quantify nonhomogeneous strain along aponeuroses.     

Passive knee movement-induced modulation of the soleus H-reflex and alteration in the fascicle length of the medial gastrocnemius muscle in humans

In humans, an inhibitory via Ia afferent pathway from the medial gastrocnemius (MG) to the soleus (SOL) motoneuron pool has been suggested. Herein, we examined the relation between MG fascicle length changes and the SOL H-reflex modulation during passive knee movement. Twelve subjects performed static and passive (5° s^?1) knee movement tasks with the ankle immobilized using an isokinetic dynamometer in sitting posture. The maximal H- and M-waves were measured at four target angles (20°, 40°, 60°, and 80° flexion from full knee extension). The MG fascicles length and velocity were measured using a B-mode ultrasonic apparatus. Results demonstrated that the SOL Hmax/Mmax; i.e., ratio of the maximal H- to M-waves, was attenuated with increasing MG fascicle length in static tasks. The SOL Hmax/Mmax at 20° was significantly attenuated compared with 60° and 80° with increasing MG fascicle length and lengthening velocity in passive knee extension. However, no significant differences in the SOL Hmax/Mmax were found across the target angles in the passive knee flexion task. In conclusion, as muscle spindles increase their discharge with lengthening fascicle velocity, but keep silent when fascicles shorten, our data suggest that lengthening the MG facilitates an inhibitory Ia pathway from MG to SOL, and modulates SOL motoneuron activity during movements.     

Medial gastrocnemius muscle fascicle active torque-length and Achilles tendon properties in young adults with spastic cerebral palsy

Individuals with spastic cerebral palsy (CP) typically experience muscle weakness. The mechanisms responsible for muscle weakness in spastic CP are complex and may be influenced by the intrinsic mechanical properties of the muscle and tendon. The purpose of this study was to investigate the medial gastrocnemius (MG) muscle fascicle active torque-length and Achilles tendon properties in young adults with spastic CP. Nine relatively high functioning young adults with spastic CP (GMFCS I, 17±2 years) and 10 typically developing individuals (18±2 years) participated in the study. Active MG torque-length and Achilles tendon properties were assessed under controlled conditions on a dynamometer. EMG was recorded from leg muscles and ultrasound was used to measure MG fascicle length and Achilles tendon length during maximal isometric contractions at five ankle angles throughout the available range of motion and during passive rotations imposed by the dynamometer. Compared to the typically developing group, the spastic CP group had 33% lower active ankle plantarflexion torque across the available range of ankle joint motion, partially explained by 37% smaller MG muscle and 4% greater antagonistic co-contraction. The Achilles tendon slack length was also 10% longer in the spastic CP group. This study confirms young adults with mild spastic CP have altered muscle–tendon mechanical properties. The adaptation of a longer Achilles tendon may facilitate a greater storage and recovery of elastic energy and partially compensate for decreased force and work production by the small muscles of the triceps surae during activities such as locomotion.     

Estimation of passive ankle joint moment during standing and walking

This study estimated the passive ankle joint moment during standing and walking initiation and its contribution to total ankle joint moment during that time. The decrement of passive joint moment due to muscle fascicle shortening upon contraction was taken into account. Muscle fascicle length in the medial gastrocnemius, which was assumed to represent muscle fascicle length in plantarflexors, was measured using ultrasonography during standing, walking initiation, and cyclical slow passive ankle joint motion. Total ankle joint moment during standing and walking initiation was calculated from ground reaction forces and joint kinematics. Passive ankle joint moment during the cyclical ankle joint motion was measured via a dynamometer. Passive ankle joint moment during standing and at the time (Tp) when the MG muscle-tendon complex length was longest in the stance phase during walking initiation were 2.3 and 5.4 Nm, respectively. The muscle fascicle shortened by 2.9 mm during standing compared with the length at rest, which decreased the contribution of passive joint moment from 19.9% to 17.4%. The muscle fascicle shortened by 4.3 mm at Tp compared with the length at rest, which decreased the contribution of passive joint moment from 8.0% to 5.8%. These findings suggest that (a) passive ankle joint moment plays an important role during standing and walking initiation even in view of the decrement of passive joint moment due to muscle fascicle shortening upon muscle contraction, and (b) muscle fascicle shortening upon muscle contraction must be taken into account when estimating passive joint moment during movements.     

Acute effect of muscle stretching on the steadiness of sustained submaximal contractions of the plantar flexor muscles

This paper examines the acute effect of a bout of static stretches on torque fluctuation during an isometric torque-matching task that required subjects to sustain isometric contractions as steady as possible with the plantar flexor muscles at four intensities (5, 10, 15, and 20% of maximum) for 20 s. The stretching bout comprised five 60-s passive stretches, separated by 10-s rest. During the torque-matching tasks and muscle stretching, the torque (active and passive) and surface electromyogram (EMG) of the medial gastrocnemius (MG), soleus (Sol), and tibialis anterior (TA) were continuously recorded. Concurrently, changes in muscle architecture (fascicle length and pennation angle) of the MG were monitored by ultrasonography. The results showed that during stretching, passive torque decreased and fascicle length increased gradually. Changes in these two parameters were significantly associated (r(2) = 0.46; P < 0.001). When data from the torque-matching tasks were collapsed across the four torque levels, stretches induced greater torque fluctuation (P < 0.001) and enhanced EMG activity (P < 0.05) in MG and TA muscles with no change in coactivation. Furthermore, stretching maneuvers produced a greater decrease (～15%; P < 0.001) in fascicle length during the torque-matching tasks and change in torque fluctuation (CV) was positively associated with changes in fascicle length (r(2) = 0.56; P < 0.001), MG and TA EMG activities, and coactivation (r(2) = 0.35, 0.34, and 0.35, respectively; P < 0.001). In conclusion, these observations indicate that repeated stretches can decrease torque steadiness by increasing muscle compliance and EMG activity of muscles around the joint. The relative influence of such adaptations, however, may depend on the torque level during the torque-matching task.     

Fascicle length of gastrocnemius muscles in monozygous twins

A large inter-individual variation is seen in muscle fascicle length of the athletes but the reasons for this phenomenon are unclear. The purpose of this study was to determine whether genetic factors contribute to the variances in muscle architectural characteristics. Nine monozygous twin pairs (3 males and 6 females), mean age 23 years (range 17-40) were studied. Fascicle length, pennation angle, and muscle thickness of the medial (MG) and lateral (LG) gastrocnemius muscles were measured in vivo by B-mode ultrasound. In the LG muscle intrapair resemblance (P < 0.01) for fascicle length (r = 0.98), pennation angle (r = 0.94) and muscle thickness (r = 0.86) were observed. In MG muscle, however, there was no intrapair resemblance for fascicle length (r = 0.66, P > 0.05), although pennation angle (r = 0.73, P < 0.05) and muscle thickness (r = 0.86, P < 0.01) were significant. Mean percent intrapair difference in LG and MG muscles were 1.8% and 5.1% for fascicle length, 11.3% and 12.3% for pennation angle and 12.4% and 9.9% for muscle thickness, respectively. There is intrapair difference between muscle thickness and pennation angle in both MG (r = 0.69, P < 0.05) and LG (r = 0.70, P < 0.05) muscles. However, no significant correlation was observed for intrapair difference between muscle thickness and fascicle length in both muscles (MG, r = 0.46; LG, r = 0.40). It appears that genetic predisposition is the predominant factor for the determination of muscle fascicle length. However, a lack of intrapair resemblance in MG fascicle length raises the possibility that fascicle length may be further influenced by external environmental factors such as physical training.     

Asymmetric deformation of contracting human gastrocnemius muscle

Muscle fiber deformation is related to its cellular structure, as well as its architectural arrangement within the musculoskeletal system. While playing an important role in aponeurosis displacement, and efficiency of force transmission to the tendon, such deformation also provides important clues about the underlying mechanical structure of the muscle. We hypothesized that muscle fiber cross section would deform asymmetrically to satisfy the observed constant volume of muscle during a contraction. Velocity-encoded, phase-contrast, and morphological magnetic resonance imaging techniques were used to measure changes in fascicle length, pinnation angle, and aponeurosis separation of the human gastrocnemius muscle during passive and active eccentric ankle joint movements. These parameters were then used to subsequently calculate the in-plane muscle area subtended by the two aponeuroses and fascicles and to calculate the in-plane (dividing area by fascicle length), and through-plane (dividing muscle volume by area) thicknesses. Constant-volume considerations of the whole-muscle geometry require that, as fascicle length increases, the muscle fiber cross-sectional area must decrease in proportion to the length change. Our empirical findings confirm the definition of a constant-volume rule that dictates that changes in the dimension perpendicular to the plane, i.e., through-plane thickness, (-6.0% for passive, -3.3% for eccentric) equate to the reciprocal of the changes in area (6.8% for passive, 3.7% for eccentric) for both exercise paradigms. The asymmetry in fascicle cross-section deformation for both passive and active muscle fibers is established in this study with a ～22% in-plane and ～6% through-plane fascicle thickness change. These fiber deformations have functional relevance, not only because they affect the force production of the muscle itself, but also because they affect the characteristics of adjacent muscles by deflecting their line of pull.     

The length dependence of muscle active force: considerations for parallel elastic properties.

The purpose of this study was to choose between two popular models of skeletal muscle: one with the parallel elastic component in parallel with both the contractile element and the series elastic component (model A), and the other in which it is in parallel with only the contractile element (model B). Passive and total forces were obtained at a variety of muscle lengths for the medial gastrocnemius muscle in anesthetized rats. Passive force was measured before the contraction (passive A) or was estimated for the fascicle length at which peak total force occurred (passive B). Fascicle length was measured with sonomicrometry. Active force was calculated by subtracting passive (A or B) force from peak total force at each fascicle or muscle length. Optimal length, that fascicle length at which active force is maximized, was 13.1 +/- 1.2 mm when passive A was subtracted and 14.0 +/- 1.1 mm with passive B (P < 0.01). Furthermore, the relationship between double-pulse contraction force and length was broader when calculated with passive B than with passive A. When the muscle was held at a long length, passive force decreased due to stress relaxation. This was accompanied by no change in fascicle length at the peak of the contraction and only a small corresponding decrease in peak total force. There is no explanation for the apparent increase in active force that would be obtained when subtracting passive A from the peak total force. Therefore, to calculate active force, it is appropriate to subtract passive force measured at the fascicle length corresponding to the length at which peak total force occurs, rather than passive force measured at the length at which the contraction begins.     

Myofascial force transmission between the human soleus and gastrocnemius muscles during passive knee motion

The plantarflexors of the lower limb are often assumed to act as independent actuators, but the validity of this assumption is the subject of considerable debate. This study aims to determine the degree to which passive changes in gastrocnemius muscle length, induced by knee motion, affect the tension in the adjacent soleus muscle. A second aim is to quantify the magnitude of myofascial passive force transmission between gastrocnemius and adjacent soleus. Fifteen healthy volunteers participated. Simultaneous ultrasound images of the gastrocnemius and soleus muscles were obtained during passive knee flexion (0-90°), while keeping the ankle angle fixed at either 70° or 115°. Image correlation analysis was used to quantify muscle fascicle lengths in both muscles. The data show that the soleus muscle fascicles elongate significantly during gastrocnemius shortening. The approximate change in passive soleus force as a result of the observed change in fascicle length was estimated and appears to be <5 N, but this estimate is sensitive to the assumed slack length of soleus.     

The morphology of the medial gastrocnemius in typically developing children and children with spastic hemiplegic cerebral palsy

We collected 3D ultrasound images of the medial gastrocnemius muscle belly (MG) in 16 children with spastic hemiplegic cerebral palsy (SHCP) (mean age: 7.8 years; range: 4–12) and 15 typically-developing (TD) children (mean age: 9.5 years; range: 4–13). All children with SHCP had limited passive dorsiflexion range on the affected side with the knee extended (mean ± 1SD: −9.3° ± 11.8). Scans were taken of both legs with the ankle joint at its resting angle (RA) and at maximum passive dorsiflexion (MD), with the knee extended. RA and MD were more plantar flexed (p < 0.05) in children with SHCP than in TD children.

We measured the volumes and lengths of the MG bellies. We also measured the length of muscle fascicles in the mid-portion of the muscle belly and the angle that the fascicles made with the deep aponeurosis of the muscle. Volumes were normalised to the subject’s body mass; muscle lengths and fascicle lengths were normalised to the length of the fibula.

Normalised MG belly lengths in the paretic limb were shorter than the non-paretic side at MD (p = 0.0001) and RA (p = 0.0236). Normalised muscle lengths of the paretic limb were shorter than those in TD children at both angles (p = 0.0004; p = 0.0003). However, normalised fascicle lengths in the non-paretic and paretic limbs were similar to those measured in TD children (p > 0.05). When compared to the non-paretic limb, muscle volume was reduced in the paretic limb (p < 0.0001), by an average of 28%, and normalised muscle volume in the paretic limb was smaller than in the TD group (p < 0.0001).

The MG is short and small in the paretic limb of children with SHCP. The altered morphology is not due to a decrease in fascicle length. We suggest that MG deformity in SHCP is caused by lack of cross-sectional growth.     

An algorithm for automated analysis of ultrasound images to measure tendon excursion in vivo

The accuracy of an algorithm for the automated tracking of tendon excursion from ultrasound images was tested in three experiments. Because the automated method could not be tested against direct measurements of tendon excursion in vivo, an indirect validation procedure was employed. In one experiment, a wire "phantom" was moved a known distance across the ultrasound probe and the automated tracking results were compared with the known distance. The excursion of the musculotendinous junction of the gastrocnemius during frontal and sagittal plane movement of the ankle was assessed in a single cadaver specimen both by manual tracking and with a cable extensometer sutured to the gastrocnemius muscle. A third experiment involved estimation of Achilles tendon excursion in vivo with both manual and automated tracking. Root mean squared (RMS) error was calculated between pairs of measurements after each test. Mean RMS errors of less than 1 mm were observed for the phantom experiments. For the in vitro experiment, mean RMS errors of 8-9% of the total tendon excursion were observed. Mean RMS errors of 6-8% of the total tendon excursion were found in vivo. The results indicate that the proposed algorithm accurately tracks Achilles tendon excursion, but further testing is necessary to determine its general applicability.     

Passive muscle mechanical properties of the medial gastrocnemius in young adults with spastic cerebral palsy

Individuals with spastic cerebral palsy (SCP) exhibit restricted joint range of motion and increased joint stiffness due to structural alterations of their muscles. Little is known about which muscle–tendon structures are responsible for these alterations. The aim of this study was to compare the passive mechanics of the ankle joint and medial gastrocnemius (MG) muscle in young adults with SCP and typically developed (TD) individuals. Nine ambulant SCP (17±2 years) and ten TD individuals (18±2 years) participated in the study. Physiological cross sectional area was estimated using freehand 3D ultrasound and found to be 37% lower in the SCP group. An isokinetic dynamometer rotated the ankle through its range while joint torque and ultrasound images of the MG muscle fascicles were simultaneously measured. Mean ankle stiffness was found to be 51% higher and mean MG fascicle strain 47% lower in the SCP group. Increased resistance to passive ankle dorsiflexion in SCP appears to be related to the inability of MG muscle fascicles to elongate with increased force.     

Ultrasound-based subject-specific parameters improve fascicle behaviour estimation in Hill-type muscle model

The estimation of muscle fascicle behaviour is decisive in a Hill-type model as they are related to muscle force by the force–length–velocity relationship and the tendon force–strain relationship. This study was aimed at investigating the influence of subject-specific tendon force–strain relationship and initial fascicle geometry (IFG) on the estimation of muscle forces and fascicle behaviour during isometric contractions. Ultrasonography was used to estimate the in vivo muscle fascicle behaviour and compare the muscle fascicle length and pennation angle estimated from the Hill-type model. The calibration–prediction process of the electromyography-driven model was performed using generic or subject-specific tendon definition with or without IFG as constraint. The combination of subject-specific tendon definition and IFG led to muscle fascicle behaviour closer to ultrasound data and significant lower forces of the ankle dorsiflexor and plantarflexor muscles compared to the other conditions. Thus, subject-specific ultrasound measurements improve the accuracy of Hill-type models on muscle fascicle behaviour.     

Reproducibility of gastrocnemius medialis muscle architecture during treadmill running

The purpose of this study was to assess the reproducibility of fascicle length (FL) and pennation angle (PA) of gastrocnemius medialis (GM) muscle during running in vivo. Twelve male recreational long distance runners (mean ± SD; age: 24 ± 3 years, mass: 76 ± 7 kg) ran on a treadmill at a speed of 3.0 m/s, wearing their own running shoes, for two different 10 min sessions that were at least 2 days apart. For each test day 10 acceptable trials were recorded. Ankle and knee joint angle data were recorded by a Vicon 624 system with three cameras operating at 120 Hz. B-mode ultrasonography was used to examine fascicle length and pennation angle of gastrocnemius medialis muscle. The ultrasound probe was firmly secured on the muscle belly using a lightweight foam fixation. The results indicated that fascicle length and pennation angle demonstrated high reproducibility values during treadmill running both for within and between test days. The root mean square scores between the repeated waveforms of pennation angle and fascicle length were small (∼2° and ∼3.5 mm, respectively). However, ∼14 trials for pennation angle and ∼9 trials for fascicle length may be required in order to record accurate data from muscle architecture parameters. In conclusion, ultrasound measurements may be highly reproducible during dynamic movements such as treadmill running, provided that a proper fixation is used in order to assure the constant location and orientation of the ultrasound probe throughout the movement.     

Gastrocnemius fascicle length changes with two-joint passive movements

Predicting muscle fascicle length changes during passive movements may lead to a better understanding of muscle function. The purpose of this study was to experimentally compare fascicle length changes in the gastrocnemius during two-joint passive movements with a previously derived kinematic model based on anatomical measures from a cadaver. The ratio of passive ankle to knee motion was manipulated to generate medial gastrocnemius fascicle elongation and lateral gastrocnemius fascicle shortening. Ultrasound images from both heads of the gastrocnemius fascicles were acquired at 10 degrees knee flexion increments and compared with this kinematic model. Our results suggest that the two-joint kinematic model from which we originally based our knee and ankle movements did not adequately reflect fascicle length changes during any of the movement conditions in this study. From our data, we propose that for every degree of ankle motion the medial and lateral gastrocnemius changes 0.42 mm and 0.96 mm, respectively, whereas changes of 0.14 mm and 0.22 mm are observed for the medial and lateral gastrocnemius, respectively, during knee movements.     

The role of the stretch reflex in the gastrocnemius muscle during human locomotion at various speeds.

In the present study, the fascicle length (L(fa)) of the human medial gastrocnemius (MG) muscle was monitored to evaluate possible input from the short-latency stretch reflex (SLR) during the stance phase of running and to examine its timing at various running speeds. Eight subjects ran at 2.0, 3.5, 5.0, and 6.5 m/s. The L(fa) was measured with the high-speed ultrasound fascicle scanning together with kinematics and myoelectrical activities. The amplitudes and onset latency of SLR activities were determined. During ground contact, the sudden MG fascicle stretch occurred during the early contact at all running speeds. This was followed by the fascicle shortening. The timing of fascicle stretch depended on running speed and type of foot contact. In slower speed conditions (2.0, 3.5, 5 m/s), the MG fascicle stretch and the corresponding SLR activities occurred during the middle of the braking phase. In fast-speed running (6.5 m/s), however, the MG fascicle stretch occurred later compared with the lower speed. The corresponding SLR activities occurred significantly later at the end of the braking phase. In addition to the clear demonstration of the different timings of SLR in MG during ground contact of running, the results imply that the role of the MG SLR during the stance phase of running can be different between fast- and slow-speed running conditions.     

In vivo fascicle behavior of synergistic muscles in concentric and eccentric plantar flexions in humans.

Ultrasonography was used to directly measure in vivo fascicle behavior of the medial gastrocnemius (MG) and soleus (SOL) muscles while the subjects (n=6 men) performed maximal voluntary concentric and eccentric plantar flexions at 60, 120, 180 and 240 deg/s. Fascicle shortening and lengthening velocities of MG, obtained from fascicle length changes over time, were significantly higher than those of SOL at +/-120, +/-180 and +240 deg/s, possibly reflecting physiological and mechanical differences between these muscles. On the other hand, the effective fascicle shortening and lengthening velocities, defined as the velocities in the longitudinal direction of muscle belly, were not significantly different between MG and SOL. This could be due to difference in fascicle architecture and/or the existence of mechanical linkages between these muscles. Moreover, when the contribution of tendinous tissues to muscle-tendon complex length change was determined from fascicle length, pennation angle, moment arm and joint angle, it accounted for approximately 50% in both concentric and eccentric trials, but showed considerable intra-subject variations. This result quantifiably demonstrates the importance of tendinous tissues in isokinetically controlled joint movements.     

Ultrasound estimates of Achilles tendon exhibit unexpected shortening during ankle plantarflexion

The purpose of this study was to investigate Achilles tendon (AT) length changes during a series of tasks that involved combinations of higher/lower force, and larger/smaller length changes of the medial gastrocnemius muscle-tendon unit (MTU). We sought to determine if common ultrasound-based estimates of AT length change were consistent with expectations for a passive elastic tendon acting in series with a muscle. We tested 8 healthy individuals during restricted joint calf contractions (high force, low displacement), ankle dorsi-/plantar-flexion (DF/PF) with the foot in the air (low force, high displacement), and heel raises (high force, high displacement). We experimentally estimated AT length change using two ultrasound methods, one based on muscle-tendon junction (MTJ) tracking and one based on muscle fascicle (MF) tracking. Estimates of AT length change were consistent with model expectations during restricted calf contractions, when the MTU underwent minimal length change. However, estimates of AT length changes were inconsistent with model expectations during the ankle DF/PF and heel raise tasks. Specifically, the AT was estimated to shorten substantially, often 10–20 mm, when the ankle plantarflexed beyond neutral position, despite loading conditions in which a passive, stiff spring would be expected to either lengthen (under increasing force) or maintain its length (under low force). These unexpected findings suggest the need for improvements in how we conceptually model and/or experimentally estimate MTU dynamics in vivo during motion analysis studies, particularly when the ankle plantarflexes beyond neutral.     

Reports of the length dependence of fatigue are greatly exaggerated.

Relative force depression associated with muscle fatigue is reported to be greater when assessed at short vs. long muscle lengths. This appears to be due to a rightward shift in the force-length relationship. This rightward shift may be caused by stretch of in-series structures, making sarcomere lengths shorter at any given muscle length. Submaximal force-length relationships (twitch, double pulse, 50 Hz) were evaluated before and after repetitive contractions (50 Hz, 300 ms, 1/s) in an in situ preparation of the rat medial gastrocnemius muscle. In some experiments, fascicle lengths were measured with sonomicrometry. Before repetitive stimulation, fascicle lengths were 11.3 +/- 0.8, 12.8 +/- 0.9, and 14.4 +/- 1.2 mm at lengths corresponding to -3.6, 0, and 3.6 mm where 0 is a reference length that corresponds with maximal active force for double-pulse stimulation. After repetitive stimulation, there was no change in fascicle lengths; these lengths were 11.4 +/- 0.8, 12.6 +/- 0.9, and 14.2 +/- 1.2 mm. The length dependence of fatigue was, therefore, not due to a stretch of in-series structures. Interestingly, the rightward shift that was evident when active force was calculated in the traditional way (subtraction of the passive force measured before contraction) was not seen when active force was calculated by subtracting the passive force that was associated with the fascicle length reached at the peak of the contraction. This calculation is based on the assumption that passive force decreases as the fascicles shorten during a fixed-end contraction. This alternative calculation revealed similar postfatigue absolute active force depression at all lengths. In relative terms, a length dependence of fatigue was still evident, but this was greatly diminished compared with that observed when active force was calculated with the traditional method.