Time dependence of series elasticity in tracheal smooth muscle

The stress-strain curve for the series elastic component (SEC) of tracheal smooth muscle was obtained by quick releasing the muscle from isometric tension to various afterloads and measuring the elastic recoils (SEC lengths) at a specific time after stimulation. A family of such curves was obtained by releasing the muscle at different points in time during contraction. Stiffnesses of the SEC (slopes of the stress-strain curves) at a specific stress level calculated from these curves (constant-stress stiffness) showed significant difference from one another. The same difference can also be characterized by the slope of the linear stiffness-stress curve, the constant A. The constant A during a 10-s isometric contraction was maximal at 2 s. It then decreased with time. This stiffness behavior is only seen when the effect of stress is held constant or eliminated. If stress is allowed to increase with time as it does during a tetanus then stiffness appears to increase monotonically. The SEC stiffness during active contraction was found to vary within the boundaries of the stiffness of muscle in rigor (upper limit) and that at resting state (lower limit).     

Force-velocity shifts with repetitive isometric and isotonic contractions of canine gastrocnemius in situ.

The force-velocity (F-V) relationships of canine gastrocnemius-plantaris muscles at optimal muscle length in situ were studied before and after 10 min of repetitive isometric or isotonic tetanic contractions induced by electrical stimulation of the sciatic nerve (200-ms trains, 50 impulses/s, 1 contraction/s). F-V relationships and maximal velocity of shortening (Vmax) were determined by curve fitting with the Hill equation. Mean Vmax before fatigue was 3.8 +/- 0.2 (SE) average fiber lengths/s; mean maximal isometric tension (Po) was 508 +/- 15 g/g. With a significant decrease of force development during isometric contractions (-27 +/- 4%, P < 0.01, n = 5), Vmax was unchanged. However, with repetitive isotonic contractions at a low load (P/Po = 0.25, n = 5), a significant decrease in Vmax was observed (-21 +/- 2%, P < 0.01), whereas Po was unchanged. Isotonic contractions at an intermediate load (P/Po = 0.5, n = 4) resulted in significant decreases in both Vmax (-26 +/- 6%, P < 0.05) and Po (-12 +/- 2%, P < 0.01). These results show that repeated contractions of canine skeletal muscle produce specific changes in the F-V relationship that are dependent on the type of contractions being performed and indicate that decreases in other contractile properties, such as velocity development and shortening, can occur independently of changes in isometric tension.     

A non-cross-bridge stiffness in activated frog muscle fibers

Force responses to fast ramp stretches of various amplitude and velocity, applied during tetanic contractions, were measured in single intact fibers from frog tibialis anterior muscle. Experiments were performed at 14 degrees C at approximately 2.1 microm sarcomere length on fibers bathed in Ringer's solution containing various concentrations of 2,3-butanedione monoxime (BDM) to greatly reduce the isometric tension. The fast tension transient produced by the stretch was followed by a period, lasting until relaxation, during which the tension remained constant to a value that greatly exceeded the isometric tension. The excess of tension was termed "static tension," and the ratio between the force and the accompanying sarcomere length change was termed "static stiffness." The static stiffness was independent of the active tension developed by the fiber, and independent of stretch amplitude and stretching velocity in the whole range tested; it increased with sarcomere length in the range 2.1-2.8 microm, to decrease again at longer lengths. Static stiffness increased well ahead of tension during the tetanus rise, and fell ahead of tension during relaxation. These results suggest that activation increased the stiffness of some sarcomeric structure(s) outside the cross-bridges.     

Mechanical transients of single toad stomach smooth muscle cells. Effects of lowering temperature and extracellular calcium

Smooth muscle's slow, economical contractions may relate to the kinetics of the crossbridge cycle. We characterized the crossbridge cycle in smooth muscle by studying tension recovery in response to a small, rapid length change (i.e., tension transients) in single smooth muscle cells from the toad stomach (Bufo marinus). To confirm that these tension transients reflect crossbridge kinetics, we examined the effect of lowering cell temperature on the tension transient time course. Once this was confirmed, cells were exposed to low extracellular calcium [( Ca2+]o) to determine whether modulation of the cell's shortening velocity by changes in [Ca2+]o reflected the calcium sensitivity of one or more steps in the crossbridge cycle. Single smooth muscle cells were tied between an ultrasensitive force transducer and length displacement device after equilibration in temperature-controlled physiological saline having either a low (0.18 mM) or normal (1.8 mM) calcium concentration. At the peak of isometric force, after electrical stimulation, small, rapid (less than or equal to 1.8% cell length in 3.6 ms) step stretches and releases were imposed. At room temperature (20 degrees C) in normal [Ca2+]o, tension recovery after the length step was described by the sum of two exponentials with rates of 40-90 s-1 for the fast phase and 2-4 s-1 for the slow phase. In normal [Ca2+]o but at low temperature (10 degrees C), the fast tension recovery phase slowed (apparent Q10 = 1.9) for both stretches and releases whereas the slow tension recovery phase for a release was only moderately affected (apparent Q10 = 1.4) while unaffected for a stretch. Dynamic stiffness was determined throughout the time course of the tension transient to help correlate the tension transient phases with specific step(s) in the crossbridge cycle. The dissociation of tension and stiffness, during the fast tension recovery phase after a release, was interpreted as evidence that this recovery phase resulted from both the transition of crossbridges from a low- to high-force producing state as well as a transient detachment of crossbridges. From the temperature studies and dynamic stiffness measurements, the slow tension recovery phase most likely reflects the overall rate of crossbridge cycling. From the tension transient studies, it appears that crossbridges cycle slower and have a longer duty cycle in smooth muscle. In low [Ca2+]o at 20 degrees C, little effect was observed on the form or time course of the tension transients.(ABSTRACT TRUNCATED AT 400 WORDS)     

Stiffness and tension during and after sudden length changes of glycerinated single insect fibrillar muscle fibres

Rapid length changes were applied (within 0.2 ms or 0.4 ms) to single isometrically contracted glycerol extracted muscle fibres of the dorsal longitudinal muscle ofLethocerus maximus suspended in an Ca²⁺ and ATP containing solution at 20–23‡ C. Force transients and the fibre stiffness were measured during and after rapid length changes. At length changesbelow 0.5% of the initial fibre length (∼ 2.4 Μm sarcomere length) the mechanical transients were characterized as follows: (1) After stretch and after release the force regains at least partly the value of tension before the length change within a quick phase of tension recovery. The quick phase induced by stretch was nearly completed within 1–2 ms. (2) A pulse in length of 1.5 ms duration, i.e., a stretch followed by a release to the initial length or a release followed by a stretch to the initial length, was applied to the fibre. The force transient induced by this procedure regains after the second length change the value of the isometric tension before the procedure. (3) The stiffness was constant during each length change of the “pulse” and was equal during the first and the second length changes. These findings are predicted by the muscle contraction model of Huxley and Simmons (1971): The identical force before and after a length pulse may indicate that the rotation of cross bridges after the first length change is followed by a rotation into the original position after the second length change. The constancy of the stiffness during the length changes may indicate a Hookean elastic element of the cross bridge. The similarity of the stiffness during the first and the second length changes, i.e., before and after the quick phase, gives evidence that the quick phases after stretch and after release are not accompanied by a change in the net number of attached cross bridges. If stretches ofmore than 0.5% of the initial length were applied, the mechanical transient of the muscle fibre changed as follows: (1) An ultra fast tension decay phase (duration < 0.4 ms) was observed in addition to the slower decay phase induced by the smaller stretches. (2) If the initial stretch was followed by a release to the initial length, no fast recovery phase was observed, which returns the force to the value before the stretch. The reduced tension value persists for a longer period in time than 10 ms. (3) If the muscle was stretched and released repetitively an ultra fast quick phase was induced only by the first stretch. (4) The stiffness increased during stretch, but was found to be the same in the isometrically contracting muscle and after the quick tension decay phase following a large stretch. These findings indicate that the contraction model of Huxley and Simmons has to be extended by a further process additional to cross bridge rotation in case of large stretches (> 0.5%L ini). The findings are taken to indicate a rapid detachment and reattachment of overstrained cross bridges, i.e., a cross bridge slippage induced by large stretches.     

Force enhancement without changes in cross-bridge turnover kinetics: the effect of EMD 57033.

The thiadiazinon derivative EMD 57033 has been found previously in cardiac muscle to increase isometric force generation without a proportional increase in fiber ATPase, thus causing a reduction in tension cost. To analyze the mechanism by which EMD 57033 affects the contractile system, we studied its effects on isometric force, isometric fiber ATPase, the rate constant of force redevelopment (k(redev)), active fiber stiffness, and its effect on Fo, which is the force contribution of a cross-bridge in the force-generating states. We used chemically skinned fibers of the rabbit psoas muscle. It was found that with 50 microM EMD 57033, isometric force increases by more than 50%, whereas Kredev, active stiffness, and isometric fiber ATPase increase by at most 10%. The results show that EMD 57033 causes no changes in cross-bridge turnover kinetics and no changes in active fiber stiffness that would result in a large enough increase in occupancy of the force-generating states to account for the increase in active force. However, plots of force versus length change recorded during stretches and releases (T plots) indicate that in the presence of EMD 57033 the y(o) value (x axis intercept) for the cross-bridges becomes more negative while its absolute value increases. This might suggest a larger cross-bridge strain as the basis for increased active force. Analysis of T plots with and without EMD 57033 shows that the increase in cross-bridge strain is not due to a redistribution of cross-bridges among different force-generating states favoring states of larger strain. Instead, it reflects an increased cross-bridge strain in the main force-generating state. The direct effect of EMD 57033 on the force contribution of cross-bridges in the force-generating states represents an alternative mechanism for a positive inotropic intervention.     

A study of the relationship between muscle length, tension-time area and stiffness in cat areflexic soleus

The relationship between muscle length and both tension-time area and stiffness were studied in the isolated cat soleus muscle during tetanic isometric contraction at different stimulus rates. The results show that: The area subtending the tension curve remains constant in a range of 8-10 mm of muscle length, for stimulation frequencies between 15 and 66 Hz. The muscle stiffness, measured using different amplitude stretches, remains constant over changes in muscle length of 10 mm. The stiffness is higher for smaller stretches than for larger ones. The data therefore show an approximately constant tension-time area and no significant changes in stiffness for variations in muscle length that exceed the physiological length variations during quiet standing. These results are discussed in the context of postural mechanisms.     

Human plantarflexor stiffness to multiple single-stretch trials.

The purpose of this investigation was to determine the influence of different stretch velocities, different rates of pre-stretch force development, and different pre-stretch muscle lengths on the intrinsic stiffness exhibited by the quasi-statically contracting active human plantarflexors during multiple single-stretch trials at 20-60% of maximum isometric contraction. Subjects were positioned prone, with the knee flexed 1.57 rad(90 degrees), shank stabilized, and foot secured in a hard plastic orthotic. Slowly increasing isometric plantarflexion force was produced until the plantarflexors were stretched by a rapid 0.2 rad (12 degrees) dorsiflexion movement. Plantarflexion forces and ankle positions were determined during these stretches as well as during resting stretches when the muscle was inactive. Resting forces were subtracted from the active trials, forces converted to torques, and stiffnesses determined for the first 62 ms of the stretch. The slope of the stiffness vs pre-stretch torque relationship averaged 4.30 +/- 0.34 Nm rad-1 Nm-1. Little difference was found between stiffness determined through the single-stretch method and the results of previous studies employing different mechanical inputs. Differences in stiffnesses with different stretching velocities were caused by computational artifact rather than by differences in intrinsic muscular reaction. Faster rates of pre-stretch force increase prior to the stretch resulted in slightly lower stiffnesses. Different pre-stretch muscle lengths apparently did not result in different stiffnesses. The shape of the torque vs displacement curve was remarkably insensitive to the planned manipulations of the testing conditions, responding in a stereotypical manner.     

Active and passive components in the length-dependent stiffness of tracheal smooth muscle during isotonic shortening.

Contraction of smooth muscle tissue involves interactions between active and passive structures within the cells and in the extracellular matrix. This study focused on a defined mechanical behavior (shortening-dependent stiffness) of canine tracheal smooth muscle tissues to evaluate active and passive contributions to tissue behavior. Two approaches were used. In one, mechanical measurements were made over a range of temperatures to identify those functions whose temperature sensitivity (Q(10)) identified them as either active or passive. Isotonic shortening velocity and rate of isometric force development had high Q(10) values (2.54 and 2.13, respectively); isometric stiffness showed Q(10) values near unity. The shape of the curve relating stiffness to isotonic shortening lengths was unchanged by temperature. In the other approach, muscle contractility was reduced by applying a sudden shortening step during the rise of isometric tension. Control contractions began with the muscle at the stepped length so that properties were measured over comparable length ranges. Under isometric conditions, redeveloped isometric force was reduced, but the ratio between force and stiffness did not change. Under isotonic conditions beginning during force redevelopment at the stepped length, initial shortening velocity and the extent of shortening were reduced, whereas the rate of relaxation was increased. The shape of the curve relating stiffness to isotonic shortening lengths was unchanged, despite the step-induced changes in muscle contractility. Both sets of findings were analyzed in the context of a quasi-structural model describing the shortening-dependent stiffness of lightly loaded tracheal muscle strips.     

High ionic strength and low pH detain activated skinned rabbit skeletal muscle crossbridges in a low force state

The effects of varying pH and ionic strength on the force-velocity relations and tension transients of skinned rabbit skeletal muscle were studied at 1-2 degrees C. Both decreasing pH from 7.35 to 6.35 and raising ionic strength from 125 to 360 mM reduced isometric force by about half and decreased sarcomere stiffness by about one-fourth, so that the stiffness/force ratio was increased by half. Lowering pH also decreased maximum shortening velocity by approximately 29%, while increasing ionic strength had little effect on velocity. These effects on velocity were correlated with asymmetrical effects on stiffness. The increase in the stiffness/force ratio with both interventions was manifest as a greater relative force change associated with a sarcomere length step. This force difference persisted for a variable time after the step. At the high ionic strength the force difference was long- lasting after stretches but relaxed quickly after releases, suggesting that the structures responsible would not impose much resistance to steady-state shortening. The opposite was found in the low pH experiments. The force difference relaxed quickly after stretches but persisted for a long time after releases. Furthermore, this force difference reached a constant value of approximately 8% of isometric force with intermediate sizes of release, and was not increased with larger releases. This value was almost identical to the value of an internal load that would be sufficient to account for the reduction in maximum velocity seen at the low pH. The results are interpreted as showing that both low pH and high ionic strength inhibit the movement of crossbridges into the force-generating parts of their cycle after they have attached to the actin filaments, with very few other effects on the cycle. The two interventions are different, however, in that detained bridges can be detached readily by shortening when the detention is caused by high ionic strength but not when it is caused by low pH.     

A reexamination of the thermoelastic effect in active striated muscle

Isolated Rana pipiens sartorius muscles at 0degreeC were stimulated via their nerves and small stretches or releases applied during the plateau of the isometric tetanus at lo. Extra heat above the isometric maintenance heat was produced during the drop in tension caused by release and, for very small releases (delta less than or equal to 0.5% lo), was completely reabsorbed during tension recovery. The extra heat was always directly proportional to the tension change. Heat absorption proportional to the tension change was also observed during the increase in tension produced by small stretches in the range 0.5% lo less than or equal to deltal less than or equal to 3.0% lo. The mean heat:tension ratio R in seven experiments was -0.0084, which is within the range of values reported previously by Woledge. In addition, it was found that during tension recovery after small releases of 1.0% lo less than or equal to deltal less than or equal to 3.0% lo the "contractile" component seems able to shorten about 1% lo without producing shortening heat.     

Characterization of cross-bridge elasticity and kinetics of cross-bridge cycling during force development in single smooth muscle cells

Force development in smooth muscle, as in skeletal muscle, is believed to reflect recruitment of force-generating myosin cross-bridges. However, little is known about the events underlying cross-bridge recruitment as the muscle cell approaches peak isometric force and then enters a period of tension maintenance. In the present studies on single smooth muscle cells isolated from the toad (Bufo marinus) stomach muscularis, active muscle stiffness, calculated from the force response to small sinusoidal length changes (0.5% cell length, 250 Hz), was utilized to estimate the relative number of attached cross-bridges. By comparing stiffness during initial force development to stiffness during force redevelopment immediately after a quick release imposed at peak force, we propose that the instantaneous active stiffness of the cell reflects both a linearly elastic cross-bridge element having 1.5 times the compliance of the cross-bridge in frog skeletal muscle and a series elastic component having an exponential length-force relationship. At the onset of force development, the ratio of stiffness to force was 2.5 times greater than at peak isometric force. These data suggest that, upon activation, cross-bridges attach in at least two states (i.e., low-force-producing and high-force-producing) and redistribute to a steady state distribution at peak isometric force. The possibility that the cross-bridge cycling rate was modulated with time was also investigated by analyzing the time course of tension recovery to small, rapid step length changes (0.5% cell length in 2.5 ms) imposed during initial force development, at peak force, and after 15 s of tension maintenance. The rate of tension recovery slowed continuously throughout force development following activation and slowed further as force was maintained. Our results suggest that the kinetics of force production in smooth muscle may involve a redistribution of cross-bridge populations between two attached states and that the average cycling rate of these cross-bridges becomes slower with time during contraction.     

The effect of tendon viscoelastic stiffness on the dynamic performance of isometric muscle

The effect of tendons viscoelastic stiffness on the dynamic response of the cat's tibialis anterior muscle under isometric conditions was determined. It was shown that the dynamic response model of the muscle derived under sinusoidal contraction-relaxation in the range of 20-80% of its maximal isometric tension was not statistically different before and after the disection of the whole distal tendon. It was suggested that under isometric conditions in the force range of 20-80% of the maximal, the tendon acts as a very stiff force transmission linkage without significantly modifying the muscle's performance.     

Electromechanical delay in human skeletal muscle under concentric and eccentric contractions

In contraction of skeletal muscle a delay exists between the onset of electrical activity and measurable tension. This delay in electromechanical coupling has been stated to be between 30 and 100 ms. Thus, in rapid movements it may be possible for electromyographic (EMG) activity to have terminated before force can be detected. This study was designed to determine the dependence of the EMG-tension delay upon selected initial conditions at the time of muscle activation. The right forearms of 14 subjects were passively oscillated by a motor-driven dynamometer through flexion-extension cycles of 135 deg at an angular velocity of approximately equal to 0.5 rad/s. Upon presentation of a visual stimulus the subjects maximally contracted the relaxed elbow flexors during flexion, extension, and under isometric conditions. The muscle length at the time of the stimulus was the same in all three conditions. An on-line computer monitoring surface EMG (Biceps and Brachioradialis) and force calculated the electromechanical delay. The mean value for the delay under eccentric condition, 49.5 ms, was significantly different (p less than 0.05) from the delays during isometric (53.9 ms) and concentric activity (55.5 ms). It is suggested that the time required to stretch the series elastic component (SEC) represents the major portion of the measured delay and that during eccentric muscle activity the SEC is in a more favorable condition for rapid force development.     

Tension recovery in permeabilized rat soleus muscle fibers after rapid shortening and restretch

Permeabilized rat soleus muscle fibers were subjected to rapid shortening/restretch protocols (20% muscle length, 20 ms duration) in solutions with pCa values ranging from 6.5 to 4.5. Force redeveloped after each restretch but temporarily exceeded the steady-state isometric tension reaching a maximum value approximately 2.5 s after relengthening. The relative size of the overshoot was <5% in pCa 6.5 and pCa 4.5 solutions but equaled 17% +/- 4% at pCa 6.0 (approximately half-maximal Ca2+ activation). Muscle stiffness was estimated during pCa 6.0 activations by imposing length steps at different time intervals after repeated shortening/restretch perturbations. Relative stiffness and relative tension were correlated (p < 0.001) during recovery, suggesting that tension overshoots reflect a temporary increase in the number of attached cross-bridges. Rates of tension recovery (k(tr)) correlated (p < 0.001) with the relative residual force prevailing immediately after restretch. Force also recovered to the isometric value more quickly at 5.7 < or = pCa < or = 5.9 than at pCa 4.5 (ANOVA, p < 0.05). These results show that k(tr) measurements underestimate the rate of isometric force development during submaximal Ca2+ activations and suggest that the rate of tension recovery is limited primarily by the availability of actin binding sites.     

The force-velocity relationship for microtubule sliding in demembranated sperm flagella of the sea urchin

We studied the relationship between the force and velocity of microtubule sliding in demembranated sperm flagella of the sea urchin, Hemicentrotus pulcherrimus, under auxotonic conditions following a quick release of the tension between sliding microtubules. The shape of the force-velocity curve was independent of the concentration of Mg-ATP over the range of 3.7 to 350 microM and appeared either linear or was the reverse of the hyperbolic curve seen for muscle. The power, calculated as the product of velocity and force, passed through a peak at c. 0.7 Fmax (the maximal isometric force). Thus, the maximal power is attained at a larger relative load than in muscle. The sliding velocity at 0.1 Fmax showed a hyperbolic dependence on Mg-ATP concentration, with a Km of 210 microM and a Vmax of 19 micron.sec-1. The maximal force did not significantly change over the Mg-ATP concentration range of 3.7 to 350 microM. These results are discussed in terms of a crossbridge model similar to the one originally proposed by Huxley. It is suggested that the dynein crossbridge cycle is characterized by a relatively rapid rate of attachment and a relatively slow rate of detachment.     

Changes in elastic characteristics of human muscle induced by eccentric exercise

The effect of an eccentric strength training programme on the muscular series elastic component (SEC) was studied on the flexors of the human elbow. The characteristics of the SEC were determined using an in situ technique derived from methods commonly used on isolated muscles. The results were expressed in terms of compliance-force and tension-extension relationships. These relationships indicate a sharp increase in compliance when tension decreases. Furthermore, for a given value of tension, the SEC compliance of the trained muscles is found to be lower than that of the untrained muscles. These results are discussed in relation to the active and passive parts of the SEC.     

Muscle performance during maximal isometric and dynamic contractions is influenced by the stiffness of the tendinous structures.

Contractile force is transmitted to the skeleton through tendons and aponeuroses, and, although it is appreciated that the mechanocharacteristics of these tissues play an important role for movement performance with respect to energy storage, the association between tendon mechanical properties and the contractile muscle output during high-force movement tasks remains elusive. The purpose of the study was to investigate the relation between the mechanical properties of the connective tissue and muscle performance in maximal isometric and dynamic muscle actions. Sixteen trained men participated in the study. The mechanical properties of the vastus lateralis tendon-aponeurosis complex were assessed by ultrasonography. Maximal isometric knee extensor force and rate of torque development (RTD) were determined. Dynamic performance was assessed by maximal squat jumps and countermovement jumps on a force plate. From the vertical ground reaction force, maximal jump height, jump power, and force-/velocity-related determinants of jump performance were obtained. RTD was positively related to the stiffness of the tendinous structures (r = 0.55, P < 0.05), indicating that tendon mechanical properties may account for up to 30% of the variance in RTD. A correlation was observed between stiffness and maximal jump height in squat jumps and countermovement jumps (r = 0.64, P < 0.05 and r = 0.55, P < 0.05). Power, force, and velocity parameters obtained during the jumps were significantly correlated to tendon stiffness. These data indicate that muscle output in high-force isometric and dynamic muscle actions is positively related to the stiffness of the tendinous structures, possibly by means of a more effective force transmission from the contractile elements to the bone.     

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.     

Recruitment,force and fatigue characteristics of quadriceps muscles of paraplegics isometrically activated by surface functional electrical stimulation

This study deals with the recruitment characteristics of unfatigued electrically stimulated quadriceps muscles of paraplegic subjects and with the time-dependent force output of these muscles under sustained stimulation conditions. Both these aspects of the performance of paralysed stimulated muscles were studied under isometric conditions and at different muscle lengths. The forces in the knee joint resulting from stimulation of the quadriceps were also calculated. Recruitment force curves due to a ramp-like stimulation function indicated a strong dependence on muscle length and demonstrated a sigmoid-shaped curve with three distinct regions: negligible force up to threshold stimulation intensity; rapid force increase; and levelling-off of the curve after which the force remains constant even though intensity is further increased. When normalized to the maximal force, recruitment was found to be independent of muscle length, generating a typical recruitment curve for every patient, under isometric stimulation. The peak forces were obtained at the same flexion angles previously published for normal subjects, but with much lower values. Muscle fatigue in tetanic isometric conditions, defined as the decrease in force due to sustained stimulation with fixed parameters, was found to be length dependent and to have a double exponential decay. The first is the acute force loss and is the more significant for functional purposes; the second is the more moderate and asymptotic region, in which partial force receovery in the form of bursts is observed.