Shortening-induced depression of voluntary force in unfatigued and fatigued human adductor pollicis muscle.

The goals of this study were to investigate adductor pollicis muscle (n = 7) force depression after maximal electrically stimulated and voluntarily activated isovelocity (19 and 306 degrees /s) shortening contractions and the effects of fatigue. After shortening contractions, redeveloped isometric force was significantly (P < 0.05) depressed relative to isometric force obtained without preceding shortening. For voluntarily and electrically stimulated contractions, relative force deficits respectively were (means +/- SE) 25.0 +/- 3.5 and 26.6 +/- 1.9% (19 degrees /s), 7.8 +/- 2.2 and 11.5 +/- 0.6% (306 degrees /s), and 23.9 +/- 4.4 and 31.6 +/- 4.7% (19 degrees /s fatigued). The relative force deficit was significantly smaller after fast compared with slow shortening contractions, whereas activation manner and fatigue did not significantly affect the deficit. It was concluded that in unfatigued and fatigued muscle the velocity-dependent relative force deficit was similar with maximal voluntary activation and electrical stimulation. These findings have important implications for experimental studies of force-velocity relationships. Moreover, if not accounted for in muscle models, they will contribute to differences observed between the predicted and the actually measured performance during in vivo locomotion.     

Catchlike property in human adductor pollicis muscle

The "catchlike" property is defined as the dramatic force increase in skeletal muscles when a single pulse is added at the onset of a sub-tetanic low-frequency stimulation train. This property has been observed in single motor units, whole animal and human muscles. It is an inherent property of muscle fibres and is not related to an increase in motor unit recruitment. Despite an abundance of observations, its origin remains unclear. The aim of this study was to induce the catchlike property in human adductor pollicis and identify its possible origin. Thumb adduction forces were measured using ulnar nerve electrical stimulation at 10Hz for reference trains (RTs) with one extra pulse 8ms after the first stimulation pulse for the experimental trains (ETs). Tests were performed at two muscle length and three stimulation levels and muscle stiffness and potentiation were quantified for all test conditions. The ETs showed higher forces and greater rates of force increase than the RTs. In addition, force increase was more pronounced at short compared to long muscle length, but no differences were found in force increase for the three stimulation levels. Furthermore, potentiation and stiffness were similar across all experimental conditions. Together, these results suggest that the increase in force associated with the catchlike property is neither caused by an increased proportion of attached cross-bridges nor potentiation of the muscle, but appears to be muscle length dependent and present in both slow and fast motor units.     

Effects of speed and distance of muscle shortening on force depression during voluntary contractions

The purpose of this study was to determine the influence of speed and distance of muscle shortening on the amount of force depression for voluntary contractions. Two experimental tests were performed. In the first test, subjects performed isometric knee extensor contractions following muscle shortening produced by isokinetic knee extensions over the range 25-50 degrees. In the second test, subjects performed isometric knee extensor contractions following muscle shortening produced by isokinetic knee extensions at two speeds: 20 and 240 degrees /s. Knee extensor moments, surface electromyographical (EMG) signals of quadriceps femoris, and interpolated twitch moments were measured during all contractions and were compared with the corresponding values obtained during purely isometric contractions. Force depression following muscle shortening for the voluntary contractions tested in this study did not depend on the distance or the speed of muscle shortening. These results are in contrast to the corresponding results in the literature obtained using artificial electrical stimulation in which force depression was always found to be directly related to the distance of shortening and inversely related to the speed of shortening. The difference in force depression as a function of the distance and speed of muscle shortening between voluntary and artificial electrical stimulation may be associated with changes in activation following the voluntary shortening contractions, whereas activation is controlled and constant in all artificial stimulation protocols.     

Stimulation frequency and force potentiation in the human adductor pollicis muscle

The effect of stimulation frequency on twitch force potentiation was examined in the adductor pollicis muscle of ten normal subjects. The ulnar nerve was supramaximally stimulated at the wrist and isometric twitch force was measured from a 3-Hz train lasting 1 s. Test stimulation frequencies of 5, 10, 20, 25, 30, 40, 50 and 100 Hz were applied for 5 s each in random order (5 min apart) and the twitches (3 Hz) were applied immediately before and after (1 s) the test frequency and at intervals up to 5 min afterwards (10 s, and 1, 2 and 5 min). Poststimulation twitches were expressed as a percentage of the prestimulation twitch. Low frequency fatigue was not induced by the protocol since the 20:50 Hz ratio did not alter within each session. The degree of twitch potentiation was frequency dependent, with potentiation increasing up to 50 Hz [mean 173 (SD 16)%] but the effect was markedly less at 100 Hz [mean 133 (SD 25)%, P less than 0.01] for all subjects. The reduced potentiation at 100 Hz may have occurred due to high frequency fatigue produced by the 100-Hz test stimulation train. The optimal frequency of those examined in the experimental group was 50 Hz but this only produced maximal potentiation in six of the ten subjects and 100 Hz always produced less potentiation. These findings have implications for electrical stimulation of muscle in the clinical setting.     

Activation-induced force enhancement in human adductor pollicis

It has been known for a long time that the steady-state isometric force after muscle stretch is bigger than the corresponding force obtained in a purely isometric contraction for electrically stimulated and maximal voluntary contractions (MVC). Recent studies using sub-maximal voluntary contractions showed that force enhancement only occurred in a sub-group of subjects suggesting that force enhancement for sub-maximal voluntary contractions has properties different from those of electrically-induced and maximal voluntary contractions. Specifically, force enhancement for sub-maximal voluntary contractions may contain an activation-dependent component that is independent of muscle stretching. To address this hypothesis, we tested for force enhancement using (i) sub-maximal electrically-induced contractions and stretch and (ii) using various activation levels preceding an isometric reference contraction at 30% of MVC (no stretch). All tests were performed on human adductor pollicis muscles. Force enhancement following stretching was found for all subjects (n = 10) and all activation levels (10%, 30%, and 60% of MVC) for electrically-induced contractions. In contrast, force enhancement at 30% of MVC, preceded by 6 s of 10%, 60%, and 100% of MVC was only found in a sub-set of the subjects and only for the 60% and 100% conditions. This result suggests that there is an activation-dependent force enhancement for some subjects for sub-maximal voluntary contractions. This activation-dependent force enhancement was always smaller than the stretch-induced force enhancement obtained at the corresponding activation levels. Active muscle stretching increased the force enhancement in all subjects, independent whether they showed activation dependence or not. It appears that post-activation potentiation, and the associated phosphorylation of the myosin light chains, might account for the stretch-independent force enhancement observed here.     

The shape of the force-elbow angle relationship for maximal voluntary contractions and sub-maximal electrically induced contractions in human elbow flexors

The force-length relationship is a basic property of skeletal muscle. Knowledge of this relationship is necessary for most analyses of human movement, and in simulation models predicting movement control strategies. Studies on animal muscles have shown that force-length relationships for sub-maximal contractions are not related through a simple scaling procedure to the relationship for maximal contractions. Furthermore, potentiation might produce a shift of sub-maximal relative to maximal force-length relationships. In this study, we tested the hypothesis that human force-elbow angle relationships for sub-maximal unpotentiated contractions are shifted to larger elbow angles (i.e. larger muscle lengths) compared to the relationship for maximal voluntary contractions (MVC), and that this shift is reduced, or even abolished, for sub-maximal potentiated contractions. Force-elbow angle relationships (48-160 degrees) were obtained from healthy subjects (n=13). At each of nine tested elbow angles, the test set consisted of a single twitch (ST(pre)) and a doublet twitch (DT(pre)) stimulation of m. biceps brachii, followed by an MVC, followed by another single twitch (ST(post)) and a doublet twitch (DT(post)) stimulation. The single and doublet twitches induced sub-maximal contractions. The force-elbow angle relationships for the pre-MVC (unpotentiated) twitch contractions were shifted to larger angles compared to those obtained for MVC. The force-elbow angle relationships for the post-MVC (potentiated) twitch contractions were shifted to smaller angles compared to those obtained for the unpotentiated twitch contractions. These results support the idea that the shift to larger muscle lengths for the sub-maximal, unpotentiated force-length relationships relative to the relationship for maximal contractions may be caused by a length-dependent Ca(2+) sensitivity that may be offset, at least in part, by potentiation.     

Postactivation potentiation in a human muscle: effect on the load-velocity relation of tetanic and voluntary shortening contractions.

Recently it was demonstrated that postactivation potentiation (PAP), which refers to the enhancement of the muscle twitch torque as a result of a prior conditioning contraction, increased the maximal rate of torque development of tetanic and voluntary isometric contractions (3). In this study, we investigated the effects of PAP and its decay over time on the load-velocity relation. To that purpose, angular velocity of thumb adduction in response to a single electrical stimulus (twitch), a high-frequency train of 15 pulses at 250 Hz (HFT(250)), and during ballistic voluntary shortening contractions, performed against loads ranging from 10 to 50% of the maximum torque, were recorded before and after a conditioning 6-s maximal voluntary contraction (MVC). The results showed an increase of the peak angular velocity for the different loads tested after the conditioning MVC (P < 0.001), but the effect was greatest for the twitch ( approximately 182%) compared with the HFT(250) or voluntary contractions ( approximately 14% for both contraction types). The maximal potentiation occurred immediately following the conditioning MVC for the twitch, whereas it was reached 1 min later for the tetanic and ballistic voluntary contractions. At that time, the load-velocity relation was significantly shifted upward, and the maximal power of the muscle was increased ( approximately 13%; P < 0.001). Furthermore, the results also indicated that the effect of PAP on shortening contractions was not related to the modality of muscle activation. In conclusion, the findings suggest a functional significance of PAP in human movements by improving muscle performance of voluntary dynamic contractions.     

Tetanic force development of adductor pollicis muscle in anesthetized man.

The tetanic force development of the human adductor pollicis muscle was studied under light anesthesia with nitrous oxide, oxygen, and Demerol, by the use of tetanic stimulation of the ulnar nerve at frequencies ranging from 10 to 100 Hz. The time necessary for the tetanic contraction to reach a plateau was longest at frequencies between 15 and 20 Hz. Fusion of tetanus occurred between 40 and 45 Hz. The mean maximal force of 6.92 kg was developed at a mean frequency of approximately 75 Hz. The maximal force was well maintained up to a stimulation frequency of 100 Hz. The results indicate that in lightly anesthetized man, the maximal force is developed at higher stimulation frequencies than those observed in conscious man and that it is well sustained at higher frequencies.     

The measurement of force/velocity relationships of fresh and fatigued human adductor pollicis muscle

The purpose of the study was to obtain force/velocity relationships for electrically stimulated (80 Hz) human adductor pollicis muscle (n = 6) and to quantify the effects of fatigue. There are two major problems of studying human muscle in situ; the first is the contribution of the series elastic component, and the second is a loss of force consequent upon the extent of loaded shortening. These problems were tackled in two ways. Records obtained from isokinetic releases from maximal isometric tetani showed a late linear phase of force decline, and this was extrapolated back to the time of release to obtain measures of instantaneous force. This method gave usable data up to velocities of shortening equivalent to approximately one-third of maximal velocity. An alternative procedure (short activation, SA) allowed the muscle to begin shortening when isometric force reached a value that could be sustained during shortening (essentially an isotonic protocol). At low velocities both protocols gave very similar data (r2 = 0.96), but for high velocities only the SA procedure could be used. Results obtained using the SA protocol in fresh muscle were compared to those for muscle that had been fatigued by 25 s of ischaemic isometric contractions, induced by electrical stimulation at the ulnar nerve. Fatigue resulted in a decrease of isometric force [to 69 (3)%], an increase in half-relaxation time [to 431 (10)%], and decreases in maximal shortening velocity [to 77 (8)%] and power [to 42 (5)%]. These are the first data for human skeletal muscle to show convincingly that during acute fatigue, power is reduced as a consequence of both the loss of force and slowing of the contractile speed.     

Behavior of human muscle fascicles during shortening and lengthening contractions in vivo.

The aim of the present study was to investigate the behavior of human muscle fascicles during dynamic contractions. Eight subjects performed maximal isometric dorsiflexion contractions at six ankle joint angles and maximal isokinetic concentric and eccentric contractions at five angular velocities. Tibialis anterior muscle architecture was measured in vivo by use of B-mode ultrasonography. During maximal isometric contraction, fascicle length was shorter and pennation angle larger compared with values at rest (P < 0.01). During isokinetic concentric contractions from 0 to 4.36 rad/s, fascicle length measured at a constant ankle joint angle increased curvilinearly from 49.5 to 69.7 mm (41%; P < 0.01), whereas pennation angle decreased curvilinearly from 14.8 to 9.8 degrees (34%; P < 0.01). During eccentric muscle actions, fascicles contracted quasi-isometrically, independent of angular velocity. The behavior of muscle fascicles during shortening contractions was believed to reflect the degree of stretch applied to the series elastic component, which decreases with increasing contraction velocity. The quasi-isometric behavior of fascicles during eccentric muscle actions suggests that the series elastic component acts as a mechanical buffer during active lengthening.     

The effect of age and gender on the relative fatigability of the human adductor pollicis muscle

The purpose of this study was to examine the relative influence of such factors as age, gender, and absolute force on the fatiguability of the human adductor pollicis muscle. 12 young males (YM, 25.3 +/- 2.1 y), 12 young females (YF. 23.5 +/- 2.1 y), 12 older males (OM, 71.7 +/- 5.6 y) and 12 older females (OF, 69.5 +/- 4.6 y) participated. Three minutes of intermittent (5 s contraction, 2 s rest) maximal voluntary contractions (MVC) were used to fatigue the adductor pollicis muscle; the ulnar nerve was also stimulated in each 2 s rest period to evoke a maximal twitch. Males were stronger than females in both voluntary and evoked force (PT) in the young age group (MVC: YM, 10.0 +/- 2.7 kg vs. YF, 6.6 +/- 1.1 kg, P < 0.05) (PT: YM, 0.99 +/- 0.21 kg vs. YF, 0.71 +/- 0.12 kg, P < 0.05). In the older adults, however, males were stronger only in the evoked twitch (OM, 0.73 +/- 0.24 kg vs. OF, 0.48 +/- 0.07 kg, P < 0.05). There was no significant effect of gender or absolute muscle force on relative fatigability; the only variable found to significantly affect fatigability was age. Older adults were significantly less fatigable than young adults as indicated by the voluntary fatigue index (FI) (percentage of force reduction from baseline; FI-young, 40.2 +/- 12.6% vs. FI-old, 25.2 +/- 12.3%). This age effect, however, was more prominent in males than females (FI-YM, 44.7 +/- 10.5% vs. FI-OM, 24.2 +/- 10.7%, P < 0.01; FI-YF, 37.8 +/- 14.1% vs. FI-OF, 26.3 +/- 14.5%, P = 0.13). In conclusion, age was found to be the strongest single predictor of fatigability during short duration, intermittent exercise in human adductor pollicis muscle.     

The relationship between force depression following shortening and mechanical work in skeletal muscle

Force depression following muscle shortening was investigated in cat soleus (n=6) at 37 degrees C for a variety of contractile conditions with the aim to test the hypotheses that force depression was independent of the speed of shortening and was directly related to the mechanical work produced by the muscle during shortening. Force depression was similar for tests in which the mechanical work performed by the muscle was similar, independent of the speed of shortening (range of speeds: 4-256mm/s). On the other hand, force depression varied significantly at a given speed of shortening but different amounts of mechanical work, supporting the hypothesis that force depression was not speed - but work dependent. The variations in the mechanical work produced by the muscle during shortening accounted for 87-96% of the variance observed in the force depression following shortening further supporting the idea that the single scalar variable work accounts for most of the observed loss in isometric force after shortening. The results of the present study are also in agreement with the notion that the mechanism underlying force depression might be associated with an inhibition of cross-bridge attachments in the overlap zone formed during the shortening phase, as proposed previously (Herzog and Leonard, 1997. Journal of Bimechanics 30 (9), 865-872; Maréchal and Plaghki, 1979.     

Repeatability of corticospinal and spinal measures during lengthening and shortening contractions in the human tibialis anterior muscle

Elements of the human central nervous system (CNS) constantly oscillate. In addition, there are also methodological factors and changes in muscle mechanics during dynamic muscle contractions that threaten the stability and consistency of transcranial magnetic stimulation (TMS) and perpherial nerve stimulation (PNS) measures.

Purpose

To determine the repeatability of TMS and PNS measures during lengthening and shortening muscle actions in the intact human tibialis anterior.

Methods

On three consecutive days, 20 males performed lengthening and shortening muscle actions at 15, 25, 50 and 80% of maximal voluntary contraction (MVC). The amplitude of the Motor Evoked Potentials (MEPs) produced by TMS was measured at rest and during muscle contraction at 90° of ankle joint position. MEPs were normalised to Mmax determined with PNS. The corticospinal silent period was recorded at 80% MVC. Hoffman reflex (H-reflex) at 10% isometric and 25% shortening and lengthening MVCs, and V-waves during MVCs were also evoked on each of the three days.

Results

With the exception of MEPs evoked at 80% shortening MVC, all TMS-derived measures showed good reliability (ICC = 0.81–0.94) from days 2 to 3. Confidence intervals (CI, 95%) were lower between days 2 and 3 when compared to days 1 and 2. MEPs significantly increased at rest from days 1 to 2 (P = 0.016) and days 1 to 3 (P = 0.046). The H-reflex during dynamic muscle contraction was reliable across the three days (ICC = 0.76–0.84). V-waves (shortening, ICC = 0.77, lengthening ICC = 0.54) and the H-reflex at 10% isometric MVC (ICC = 0.66) was generally less reliable over the three days.

Conclusion

Although it is well known that measures of the intact human CNS exhibit moment-to-moment fluctuations, careful experimental arrangements make it possible to obtain consistent and repeatable measurements of corticospinal and spinal excitability in the actively lengthening and shortening human TA muscle.     

Force recovery after activated shortening in whole skeletal muscle: transient and steady-state aspects of force depression.

The depression of isometric force after active shortening is a well-accepted characteristic of skeletal muscle, yet its mechanisms remain unknown. Although traditionally analyzed at steady state, transient phenomena caused, at least in part, by cross-bridge kinetics may provide novel insight into the mechanisms associated with force depression (FD). To identify the transient aspects of FD and its relation to shortening speed, shortening amplitude, and muscle mechanical work, in situ experiments were conducted in soleus muscle-tendon units of anesthetized cats. The period immediately after shortening, in which force recovers toward steady state, was fit by using an exponential recovery function (R2 > 0.99). Statistical analyses revealed that steady-state FD (FD(ss)) increased with shortening amplitude and mechanical work. This FD(ss) increase was always accompanied by a significant decrease in force recovery rate. Furthermore, a significant reduction in stiffness was observed after all activated shortenings, presumably because of a reduced proportion of attached cross bridges. These results were interpreted with respect to the two most prominent proposed mechanisms of force depression: sarcomere length nonuniformity theory (7, 32) and a stress-induced inhibition of cross-bridge binding in the newly formed actin-myosin overlap zone (14, 28). We hypothesized that the latter could describe both steady-state and transient aspects of FD using a single scalar variable, the mechanical work done during shortening. As either excursion (overlap) or force (stress) is increased, mechanical work increases, and cross-bridge attachment would become more inhibited, as supported by this study in which an increase in mechanical work resulted in a slower recovery to a more depressed steady-state force.     

Force depression decays during shortening in the medial gastrocnemius of the rat

Force depression due to shortening of activated skeletal muscles has previously been described to be long lasting during isometric contractions following the shortening. In the present study, using the medial gastrocnemius of the rat, effects of force depression have been made apparent during shortening by computationally partially compensating for the direct effect of shortening velocity due to the tension–velocity relation. Evidence was found for the decay and complete disappearance of force depression already during continuation of the shortening contraction to short muscle lengths.     

Myofibrillar and collagen protein synthesis in human skeletal muscle in young men after maximal shortening and lengthening contractions

We aimed to determine whether there were differences in the extent and time course of skeletal muscle myofibrillar protein synthesis (MPS) and muscle collagen protein synthesis (CPS) in human skeletal muscle in an 8.5-h period after bouts of maximal muscle shortening (SC; average peak torque = 225 +/- 7 N.m, means +/- SE) or lengthening contractions (LC; average peak torque = 299 +/- 18 N.m) with equivalent work performed in each mode. Eight healthy young men (21.9 +/- 0.6 yr, body mass index 24.9 +/- 1.3 kg/m2) performed 6 sets of 10 maximal unilateral LC of the knee extensors on an isokinetic dynamometer. With the contralateral leg, they then performed 6 sets of maximal unilateral SC with work matched to the total work performed during LC (10.9 +/- 0.7 vs. 10.9 +/- 0.8 kJ, P = 0.83). After exercise, the participants consumed small intermittent meals to provide 0.1 g.kg(-1).h(-1) of protein and carbohydrate. Prior exercise elevated MPS above rest in both conditions, but there was a more rapid rise after LC (P < 0.01). The increases (P < 0.001) in CPS above rest were identical for both SC and LC and likely represent a remodeling of the myofibrillar basement membrane. Therefore, a more rapid rise in MPS after maximal LC could translate into greater protein accretion and muscle hypertrophy during chronic resistance training utilizing maximal LC.     

Ia-afferent input to motoneurons during shortening and lengthening muscle contractions in humans.

The central nervous system employs different strategies to execute specific motor tasks. Because afferent feedback during shortening and lengthening muscle contractions differs, the neural strategy underlying these tasks may be quite distinct. Cortical drive may be adjusted or afferent input regulated. The exact mechanisms are not clear. Here, we examine the control of synaptic transmission across the Ia synapse during shortening and lengthening muscle contractions. Subjects were instructed to maintain isolated activity in a single tibialis anterior (TA) motor unit while muscle length was varied from flexion to extension and back. At a fixed interval after a firing of the active motor unit, a single electrical stimulus was applied to the common peroneal nerve to activate Ia afferents from the TA muscle. We investigated the stimulus-induced change in firing probability of 19 individual low-threshold TA motor units during shortening and lengthening contractions. Any change in firing probability depends on both pre- and postsynaptic mechanisms. In this experiment, motoneuron firing rate was similar during both contraction types. There was no difference in the firing probability between shortening and lengthening contractions (0.23 +/- 0.03 and 0.20 +/- 0.02, respectively). We suggest that there is no contraction type-specific control of Ia input to the motoneurons during shortening and lengthening muscle contractions. Cortical adjustments may have occurred.