Reconstruction of the human gastrocnemius force-length curve in vivo: part 2-experimental results

For a physiologically realistic range of joint motion and therefore range of muscle fiber lengths, only part of the force-length curve can be used in vivo; i.e., the section of the force-length curve that is expressed can vary. The purpose of this study was to determine the expressed section of the force-length relationship of the gastrocnemius for humans. Fourteen male and fourteen female subjects aged 18-27 performed maximal isometric plantar flexions in a Biodex dynamometer. Plantar flexion moments were recorded at five ankle angles:-15 degrees , 0 degrees , 15 degrees , 30 degrees , and 40 degrees , with negative angles defined as dorsiflexion. These measurements were repeated for four randomly ordered knee angles over two testing sessions 4 to 10 days apart. The algorithm of Herzog and ter Keurs (1988a) was used to reconstruct the force-length curves of the biarticular gastrocnemius. Twenty-four subjects operated over the ascending limb, three operated over the descending limb, and one operated over the plateau region. The variation found suggests that large subject groups should be used to determine the extent of normal in vivo variability in this muscle property. The possible source of the variability is discussed in terms of parameters typically used in muscle models.     

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.     

The expression of the skeletal muscle force-length relationship in vivo: A simulation study

The force-length relationship is one of the most important mechanical characteristics of skeletal muscle in humans and animals. For a physiologically realistic joint range of motion and therefore range of muscle fibre lengths only part of the force-length curve may be used in vivo, i.e. only a section of the force-length curve is expressed. A generalised model of a mono-articular muscle-tendon complex was used to examine the effect of various muscle architecture parameters on the expressed section of the force-length relationship for a 90° joint range of motion. The parameters investigated were: the ratio of tendon resting length to muscle fibre optimum length (L_TR:L_F·OPT) (varied from 0.5 to 11.5), the ratio of muscle fibre optimum length to average moment arm (L_F·OPT:r) (varied from 0.5 to 5), the normalised tendon strain at maximum isometric force (c) (varied from 0 to 0.08), the muscle fibre pennation angle (θ) (varied from 0° to 45°) and the joint angle at which the optimum muscle fibre length occurred (φ). The range of values chosen for each parameter was based on values reported in the literature for five human mono-articular muscles with different functional roles. The ratios L_TR:L_F·OPT and L_F·OPT:r were important in determining the amount of variability in the expressed section of the force-length relationship. The modelled muscle operated over only one limb at intermediate values of these two ratios (L_TR:L_F·OPT=5; L_F·OPT:r=3), whether this was the ascending or descending limb was determined by the precise values of the other parameters. It was concluded that inter-individual variability in the expressed section of the force-length relationship is possible, particularly for muscles with intermediate values of L_TR:L_F·OPT and L_F·OPT:r such as the brachialis and vastus lateralis. Understanding the potential for inter-individual variability in the expressed section is important when using muscle models to simulate movement.     

Length dependence of active force production in skeletal muscle.

The sliding filament and cross-bridge theories of muscle contraction provide discrete predictions of the tetanic force-length relationship of skeletal muscle that have been tested experimentally. The active force generated by a maximally activated single fiber (with sarcomere length control) is maximal when the filament overlap is optimized and is proportionally decreased when overlap is diminished. The force-length relationship is a static property of skeletal muscle and, therefore, it does not predict the consequences of dynamic contractions. Changes in sarcomere length during muscle contraction result in modulation of the active force that is not necessarily predicted by the cross-bridge theory. The results of in vivo studies of the force-length relationship suggest that muscles that operate on the ascending limb of the force-length relationship typically function in stretch-shortening cycle contractions, and muscles that operate on the descending limb typically function in shorten-stretch cycle contractions. The joint moments produced by a muscle depend on the moment arm and the sarcomere length of the muscle. Moment arm magnitude also affects the excursion (length change) of a muscle for a given change in joint angle, and the number of sarcomeres arranged in series within a muscle fiber determines the sarcomere length change associated with a given excursion.     

Fibre operating lengths of human lower limb muscles during walking

Muscles actuate movement by generating forces. The forces generated by muscles are highly dependent on their fibre lengths, yet it is difficult to measure the lengths over which muscle fibres operate during movement. We combined experimental measurements of joint angles and muscle activation patterns during walking with a musculoskeletal model that captures the relationships between muscle fibre lengths, joint angles and muscle activations for muscles of the lower limb. We used this musculoskeletal model to produce a simulation of muscle-tendon dynamics during walking and calculated fibre operating lengths (i.e. the length of muscle fibres relative to their optimal fibre length) for 17 lower limb muscles. Our results indicate that when musculotendon compliance is low, the muscle fibre operating length is determined predominantly by the joint angles and muscle moment arms. If musculotendon compliance is high, muscle fibre operating length is more dependent on activation level and force-length-velocity effects. We found that muscles operate on multiple limbs of the force-length curve (i.e. ascending, plateau and descending limbs) during the gait cycle, but are active within a smaller portion of their total operating range.     

Sensitivity of model predictions of muscle function to changes in moment arms and muscle-tendon properties: a Monte-Carlo analysis

Hill-type muscle models are commonly used in musculoskeletal models to estimate muscle forces during human movement. However, the sensitivity of model predictions of muscle function to changes in muscle moment arms and muscle-tendon properties is not well understood. In the present study, a three-dimensional muscle-actuated model of the body was used to evaluate the sensitivity of the function of the major lower limb muscles in accelerating the whole-body center of mass during gait. Monte-Carlo analyses were used to quantify the effects of entire distributions of perturbations in the moment arms and architectural properties of muscles. In most cases, varying the moment arm and architectural properties of a muscle affected the torque generated by that muscle about the joint(s) it spanned as well as the torques generated by adjacent muscles. Muscle function was most sensitive to changes in tendon slack length and least sensitive to changes in muscle moment arm. However, the sensitivity of muscle function to changes in moment arms and architectural properties was highly muscle-specific; muscle function was most sensitive in the cases of gastrocnemius and rectus femoris and insensitive in the cases of hamstrings and the medial sub-region of gluteus maximus. The sensitivity of a muscle's function was influenced by the magnitude of the muscle's force as well as the operating region of the muscle on its force-length curve. These findings have implications for the development of subject-specific models of the human musculoskeletal system.     

Characterization of passive elastic properties of the human medial gastrocnemius muscle belly using supersonic shear imaging

The passive elastic properties of a muscle-tendon complex are usually estimated from the relationship between the joint angle and the passive resistive torque, although the properties of the different structures crossing the joint cannot be easily assessed. This study aimed to determine the passive mechanical properties of the gastrocnemius medialis muscle (GM) using supersonic shear imaging (SSI) that allows the measurement of localized muscle shear modulus (μ). The SSI of the GM was taken for 7 subjects during passive ankle dorsiflexion at a range of knee positions performed on an isokinetic dynamometer. The relationship between normalized μ and the length of the gastrocnemius muscle-tendon units (GMTU) was very well fitted to an exponential model (0.944相似文献   

Force-length properties and functional demands of cat gastrocnemius, soleus and plantaris muscles.

The purpose of this study was to measure isometric force-length properties of cat soleus, gastrocnemius and plantaris muscle-tendon units, and to relate these properties to the functional demands of these muscles during everyday locomotor activities. Isometric force-length properties were determined using an in situ preparation, where forces were measured using buckle-type tendon transducers, and muscle-tendon unit lengths were quantified through ankle and knee joint configurations. Functional demands of the muscles were assessed using direct muscle force measurements in freely moving animals. Force-length properties and functional demands were determined for soleus, gastrocnemius and plantaris muscles simultaneously in each animal. The results suggest that isometric force-length properties of cat soleus, gastrocnemius and plantaris muscles, as well as the region of the force-length relation that is used during everyday locomotor tasks, match the functional demands.     

Influence of different shortening velocities preceding stretch on human triceps surae moment generation in vivo

The purpose of this study was to examine the influence of different shortening velocities preceding the stretch on moment generation of the triceps surae muscles and architecture of the m. gastrocnemius medialis after shortening-stretch cycles of equal magnitude in vivo. Eleven male subjects (31.6+/-5.8 years, 178.4+/-7.3cm, 80.6+/-9.6kg) performed a series of electro-stimulated (85Hz) shortening-stretch plantar flexion contractions. The shortening-stretch cycles were performed at three constant angular velocities (25, 50, 100 degrees /s) in the plantar flexion direction (shortening) and at 50 degrees /s in the dorsiflexion direction (stretching). The resultant ankle joint moments were calculated through inverse dynamics. Pennation angle and fascicle length of the m. gastrocnemius medialis at rest and during contractions were measured using ultrasonography. The corresponding ankle moments, kinematics and changes in muscle architecture were analysed at seven time intervals. An analysis of variance for repeated measurements and post hoc test with Bonferroni correction was used to check the velocity-related effects on moment enhancement (alpha=0.05). The results show an increase in pennation angles and a decrease in fascicle lengths after the shortening-stretch cycle. The ankle joint moment ratio (post to pre) was higher (p<0.01) than 1.0 indicating a moment enhancement after the shortening-stretch cycle. The found ankle joint moment enhancement was 2-5% after the shortening-stretch cycle and was independed of the shortening velocity. Furthermore, the decrease in fascicle length after the shortening-stretch cycle indicates that the moment enhancement found in the present study is underestimated at least by 1-3%. Considering that the experiments have been done at the ascending limb of the force-length curve and that force enhancement is higher at the descending and the plateau region of the force-length curve, we conclude that the moment enhancement after shortening-stretch cycle can have important physiological affects while locomotion.     

Computational modelling of muscle fibre operating ranges in the hindlimb of a small ground bird (Eudromia elegans), with implications for modelling locomotion in extinct species

The arrangement and physiology of muscle fibres can strongly influence musculoskeletal function and whole-organismal performance. However, experimental investigation of muscle function during in vivo activity is typically limited to relatively few muscles in a given system. Computational models and simulations of the musculoskeletal system can partly overcome these limitations, by exploring the dynamics of muscles, tendons and other tissues in a robust and quantitative fashion. Here, a high-fidelity, 26-degree-of-freedom musculoskeletal model was developed of the hindlimb of a small ground bird, the elegant-crested tinamou (Eudromia elegans, ~550 g), including all the major muscles of the limb (36 actuators per leg). The model was integrated with biplanar fluoroscopy (XROMM) and forceplate data for walking and running, where dynamic optimization was used to estimate muscle excitations and fibre length changes throughout both gaits. Following this, a series of static simulations over the total range of physiological limb postures were performed, to circumscribe the bounds of possible variation in fibre length. During gait, fibre lengths for all muscles remained between 0.5 to 1.21 times optimal fibre length, but operated mostly on the ascending limb and plateau of the active force-length curve, a result that parallels previous experimental findings for birds, humans and other species. However, the ranges of fibre length varied considerably among individual muscles, especially when considered across the total possible range of joint excursion. Net length change of muscle–tendon units was mostly less than optimal fibre length, sometimes markedly so, suggesting that approaches that use muscle–tendon length change to estimate optimal fibre length in extinct species are likely underestimating this important parameter for many muscles. The results of this study clarify and broaden understanding of muscle function in extant animals, and can help refine approaches used to study extinct species.     

Measuring muscle and joint geometry parameters of a shoulder for modeling purposes.

An extensive set of muscle and joint geometry parameters was measured of the right shoulder of an embalmed male. For all muscles the optimal muscle fiber length was determined by laser diffraction measurements of sarcomere length. In addition, tendon length and physiological cross-sectional area were determined. The parameter set was needed to enhance the reliability of a computer model of the shoulder (Van der Helm, 1994a,b Journal of Biomechanics 27, 527-550, 551-569). With the model, an abduction of the arm was simulated in seven positions, at 30 degrees intervals. In each of the simulated arm positions, actual sarcomere lengths were calculated from the lengths of 104 muscle elements, distributed over 16 shoulder muscles. For most muscle elements, the simulated abduction appeared to take place within the sarcomere length range in which the muscle elements can exert force. The muscle elements can then act on the ascending limb as well as on the plateau and on the descending limb of the relative force-length curves of sarcomeres. The produced data set is not only important for the refinement of shoulder modeling, but also for functional analyses of shoulder movements in general.     

Knee and ankle joint torque-angle relationships of multi-joint leg extension

The force-length-relation (F-l-r) is an important property of skeletal muscle to characterise its function, whereas for in vivo human muscles, torque-angle relationships (T-a-r) represent the maximum muscular capacity as a function of joint angle. However, since in vivo force/torque-length data is only available for rotational single-joint movements the purpose of the present study was to identify torque-angle-relationships for multi-joint leg extension. Therefore, inverse dynamics served for calculation of ankle and knee joint torques of 18 male subjects when performing maximum voluntary isometric contractions in a seated leg press. Measurements in increments of 10° knee angle from 30° to 100° knee flexion resulted in eight discrete angle configurations of hip, knee and ankle joints. For the knee joint we found an ascending-descending T-a-r with a maximum torque of 289.5° ± 43.3 Nm, which closely matches literature data from rotational knee extension. In comparison to literature we observed a shift of optimum knee angle towards knee extension. In contrast, the T-a-r of the ankle joint vastly differed from relationships obtained for isolated plantar flexion. For the ankle T-a-r derived from multi-joint leg extension subjects operated over different sections of the force-length curve, but the ankle T-a-r derived from isolated joint efforts was over the ascending limb for all subjects. Moreover, mean maximum torque of 234.7 ± 56.6 Nm exceeded maximal strength of isolated plantar flexion (185.7 ± 27.8 Nm). From these findings we conclude that muscle function between isolated and more physiological multi-joint tasks differs. This should be considered for ergonomic and sports optimisation as well as for modelling and simulation of human movement.     

Sarcomere length organization as a design for cooperative function amongst all lumbar spine muscles

The functional design of spine muscles in part dictates their role in moving, loading, and stabilizing the lumbar spine. There have been numerous studies that have examined the isolated properties of these individual muscles. Understanding how these muscles interact and work together, necessary for the prediction of muscle function, spine loading, and stability, is lacking. The objective of this study was to measure sarcomere lengths of lumbar muscles in a neutral cadaveric position and predict the sarcomere operating ranges of these muscles throughout full ranges of spine movements. Sarcomere lengths of seven lumbar muscles in each of seven cadaveric donors were measured using laser diffraction. Using published anatomical coordinate data, superior muscle attachment sites were rotated about each intervertebral joint and the total change in muscle length was used to predict sarcomere length operating ranges. The extensor muscles had short sarcomere lengths in a neutral spine posture and there were no statistically significant differences between extensor muscles. The quadratus lumborum was the only muscle with sarcomere lengths that were optimal for force production in a neutral spine position, and the psoas muscles had the longest lengths in this position. During modeled flexion the extensor, quadratus lumborum, and intertransversarii muscles lengthened so that all muscles operated in the approximate same location on the descending limb of the force-length relationship. The intrinsic properties of lumbar muscles are designed to complement each other. The extensor muscles are all designed to produce maximum force in a mid-flexed posture, and all muscles are designed to operate at similar locations of the force-length relationship at full spine flexion.     

Residual force enhancement during voluntary contractions of knee extensors and flexors at short and long muscle lengths

Residual force enhancement (RFE) is a term describing the observation that muscle tension during a contraction that includes a stretch and hold remains above that during an isometric contraction at the hold length. RFE has been observed during in vitro and in vivo experiments, but results involving voluntary contractions are mixed, particularly with respect to large muscles. The purpose of this study was to determine if RFE can be observed in large muscles such as knee extensors and flexors at joint configurations corresponding to the ascending and descending limbs of the muscle force-length curve. Two groups of twenty participants (ten males and ten females per group) performed maximum voluntary contractions on a Biodex machine in purely isometric conditions and in isometric conditions immediately following eccentric stretch. Knee extension trials were performed at 40° (short muscles) and 100° (long muscles) flexion from full extension (0°), and knee flexion trials were performed at 70° (short muscles) and 10° (long muscles) flexion. Stretch-isometric trials terminated at these angles following 30° of eccentric motion at 30°/s. Statistically-significant RFE was observed for both tasks at long-muscle joint configurations, but was not observed for either task at short-muscle joint configurations. Passive torque enhancement was also observed following muscle relaxation at long-muscle joint configurations for both tasks, but for only knee flexion at short-muscle joint configurations. These results reinforce for voluntary contractions of large muscles the RFE behavior observed in smaller muscles, and provide further evidence that RFE occurs primarily on the descending limb of the muscle force-length curve.     

Considerations on the history dependence of muscle contraction.

When a skeletal muscle that is actively producing force is shortened or stretched, the resulting steady-state isometric force after the dynamic phase is smaller or greater, respectively, than the purely isometric force obtained at the corresponding final length. The cross-bridge model of muscle contraction does not readily explain this history dependence of force production. The most accepted proposal to explain both, force depression after shortening and force enhancement after stretch, is a nonuniform behavior of sarcomeres that develops during and after length changes. This hypothesis is based on the idea of instability of sarcomere lengths on the descending limb of the force-length relationship. However, recent evidence suggests that skeletal muscles may be stable over the entire range of active force production, including the descending limb of the force-length relationship. The purpose of this review was to critically evaluate hypotheses aimed at explaining the history dependence of force production and to provide some novel insight into the possible mechanisms underlying these phenomena. It is concluded that the sarcomere nonuniformity hypothesis cannot always explain the total force enhancement observed after stretch and likely does not cause all of the force depression after shortening. There is evidence that force depression after shortening is associated with a reduction in the proportion of attached cross bridges, which, in turn, might be related to a stress-induced inhibition of cross-bridge attachment in the myofilament overlap zone. Furthermore, we suggest that force enhancement is not associated with instability of sarcomeres on the descending limb of the force-length relationship and that force enhancement has an active and a passive component. Force depression after shortening and force enhancement after stretch are likely to have different origins.     

Residual force enhancement after lengthening is present during submaximal plantar flexion and dorsiflexion actions in humans.

Stretch of an activated muscle causes a transient increase in force during the stretch and a sustained, residual force enhancement (RFE) after the stretch. The purpose of this study was to determine whether RFE is present in human muscles under physiologically relevant conditions (i.e., when stretches were applied within the working range of large postural leg muscles and under submaximal voluntary activation). Submaximal voluntary plantar flexion (PF(v)) and dorsiflexion (DF(v)) activation was maintained by providing direct visual feedback of the EMG from soleus or tibialis anterior, respectively. RFE was also examined during electrical stimulation of the plantar flexion muscles (PF(s)). Constant-velocity stretches (15 degrees /s) were applied through a range of motion of 15 degrees using a custom-built ankle torque motor. The muscles remained active throughout the stretch and for at least 10 s after the stretch. In all three activation conditions, the stable joint torque measured 9-10 s after the stretch was greater than the isometric joint torque at the final joint angle. When expressed as a percentage of the isometric torque, RFE values were 7, 13, and 12% for PF(v), PF(s), DF(v), respectively. These findings indicate that RFE is a characteristic of human skeletal muscle and can be observed during submaximal (25%) voluntary activation when stretches are applied on the ascending limb of the force-length curve. Although the underlying mechanisms are unclear, it appears that sarcomere popping and passive force enhancement are insufficient to explain the presence of RFE in these experiments.     

The effects of muscle stretching and shortening on isometric forces on the descending limb of the force-length relationship

The purpose of this study was to examine the effects of stretching and shortening on the isometric forces at different lengths on the descending limb of the force-length relationship. Cat soleus (N = 10) was stretched and shortened by various amounts on the descending limb of the force-length relationship, and the steady-state forces following these dynamic contractions were compared to the isometric forces at the corresponding muscle lengths. We found a shift of the force-length relationship to greater force values following muscle stretching, and to smaller force values following muscle shortening. Shifts in both directions critically depended on the magnitude of stretching/shortening and the final muscle length. We confirm recent findings that the steady-state isometric force following some stretch conditions clearly exceeded the maximal isometric forces at optimum muscle length, and that force enhancement was associated with an increase in the passive force, i.e., a passive force enhancement. When the passive force enhancement was subtracted from the total force enhancement, forces following stretch were always equal to or smaller than the isometric force at optimum muscle length. Together, these findings led to the conclusions: (a). that force enhancement is composed of an "active and a "passive" component; (b). that the "passive" component of force enhancement allows for forces greater than the maximal isometric forces at the muscle's optimum length; and (c). that force enhancement and force depression are critically affected by muscle length and stretch/shortening amplitude.     

Three-Dimensional Muscle Architecture and Comprehensive Dynamic Properties of Rabbit Gastrocnemius,Plantaris and Soleus: Input for Simulation Studies

The vastly increasing number of neuro-muscular simulation studies (with increasing numbers of muscles used per simulation) is in sharp contrast to a narrow database of necessary muscle parameters. Simulation results depend heavily on rough parameter estimates often obtained by scaling of one muscle parameter set. However, in vivo muscles differ in their individual properties and architecture. Here we provide a comprehensive dataset of dynamic (n = 6 per muscle) and geometric (three-dimensional architecture, n = 3 per muscle) muscle properties of the rabbit calf muscles gastrocnemius, plantaris, and soleus. For completeness we provide the dynamic muscle properties for further important shank muscles (flexor digitorum longus, extensor digitorum longus, and tibialis anterior; n = 1 per muscle). Maximum shortening velocity (normalized to optimal fiber length) of the gastrocnemius is about twice that of soleus, while plantaris showed an intermediate value. The force-velocity relation is similar for gastrocnemius and plantaris but is much more bent for the soleus. Although the muscles vary greatly in their three-dimensional architecture their mean pennation angle and normalized force-length relationships are almost similar. Forces of the muscles were enhanced in the isometric phase following stretching and were depressed following shortening compared to the corresponding isometric forces. While the enhancement was independent of the ramp velocity, the depression was inversely related to the ramp velocity. The lowest effect strength for soleus supports the idea that these effects adapt to muscle function. The careful acquisition of typical dynamical parameters (e.g. force-length and force-velocity relations, force elongation relations of passive components), enhancement and depression effects, and 3D muscle architecture of calf muscles provides valuable comprehensive datasets for e.g. simulations with neuro-muscular models, development of more realistic muscle models, or simulation of muscle packages.     

The force-length relationship of a muscle-tendon complex: experimental results and model calculations

Models are useful when studying how architectural and physiological properties of muscle-tendon complexes are related to function, because they allow for the simulation of the behaviour of such complexes during natural movements. In the construction of these models, evaluation of their accuracy is an important step. In the present study, a model was constructed to calculate the isometric force-length relationship of the rat extensor digitorum longus muscle-tendon complex. The model is based on the assumption that a muscle-tendon complex is a collection of independent units, each consisting of a muscle fibre in series with a tendon fibre. By intention, values for model parameters were derived indirectly, using only the measured maximal isometric tetanic force, the distance between origin and insertion at which it occurred (optimum lOI) and an estimate of muscle fibre optimal length. The accuracy of the calculated force-length relationship was subsequently evaluated by comparing it to the relationship measured in isometric tetanic contractions of a real complex in the rat. When the length of distal muscle fibres, measured during isometric contraction at optimal lOI of the whole complex, was used as an estimate for muscle fibre optimal length of all muscle fibre-tendon fibre units in the model, the calculated relationship was too narrow. That is, both on the ascending limb and on the descending limb the calculated tetanic force was lower than the measured tetanic force.(ABSTRACT TRUNCATED AT 250 WORDS)     

Stretch-induced,steady-state force enhancement in single skeletal muscle fibers exceeds the isometric force at optimum fiber length

Stretch-induced force enhancement has been observed in a variety of muscle preparations and on structural levels ranging from single fibers to in vivo human muscles. It is a well-accepted property of skeletal muscle. However, the mechanism causing force enhancement has not been elucidated, although the sarcomere-length non-uniformity theory has received wide support. The purpose of this paper was to re-investigate stretch-induced force enhancement in frog single fibers by testing specific hypotheses arising from the sarcomere-length non-uniformity theory. Single fibers dissected from frog tibialis anterior (TA) and lumbricals (n=12 and 22, respectively) were mounted in an experimental chamber with physiological Ringer's solution (pH=7.5) between a force transducer and a servomotor length controller. The tetantic force-length relationship was determined. Isometric reference forces were determined at optimum length (corresponding to the maximal, active, isometric force), and at the initial and final lengths of the stretch experiments. Stretch experiments were performed on the descending limb of the force-length relationship after maximal tetanic force was reached. Stretches of 2.5-10% (TA) and 5-15% lumbricals of fiber length were performed at 0.1-1.5 fiber lengths/s. The stretch-induced, steady-state, active isometric force was always equal or greater than the purely isometric force at the muscle length from which the stretch was initiated. Moreover, for stretches of 5% fiber length or greater, and initiated near the optimum length of the fiber, the stretch-enhanced active force always exceeded the maximal active isometric force at optimum length. Finally, we observed a stretch-induced enhancement of passive force. We conclude from these results that the sarcomere length non-uniformity theory alone cannot explain the observed force enhancement, and that part of the force enhancement is associated with a passive force that is substantially greater after active compared to passive muscle stretch.