We hypothesised that experienced runners would select a stride frequency closer to the optimum (minimal energy costs) than would novice runners. In addition, we expected that optimal stride frequency could simply be determined by monitoring heart rate without measuring oxygen consumption (VO2). Ten healthy males (mean ±s: 24±2 year) with no running training experience and 10 trained runners of similar age ran at constant treadmill speed corresponding to 80% of individual ventilatory threshold. For two days, they ran at seven different stride frequencies (self-selected stride frequency ±18%) imposed by a metronome. Optimal stride frequency was based on the minimum of a second-order polynomial equation fitted through steady state VO2 at each stride frequency. Running cost (mean±s) at optimal stride frequency was higher (P < 0.05) in novice (236±31 ml O2·kg(-1.) km(-1)) than trained (189±13 ml O2·kg(-1.) km(-1)) runners. Self-selected stride frequency (mean ±s; strides(.)min(-1)) for novice (77.8±2.8) and trained runners (84.4±5.3) were lower (P < 0.05) than optimal stride frequency (respectively, 84.9±5.0 and 87.1±4.8). The difference between self-selected and optimal stride frequency was smaller (P < 0.05) for trained runners. In both the groups optimal stride frequency established with heart rate was not different (P > 0.3) from optimal stride frequency based on VO2. In each group and despite limited variation between participants, optimal stride frequencies derived from VO2 and heart rate were related (r > 0.7; P < 0.05). In conclusion, trained runners chose a stride frequency closer to the optimum for energy expenditure than novices. Heart rate could be used to establish optimal stride frequency.
The effects of high-frequency initial pulses (HFIP) and posttetanic potentiation on mechanical power output during concentric contractions were examined in the in situ medial gastrocnemius of the rat with an intact origin on the femur and blood supply. Stimulation of the muscle was performed via the severed sciatic nerve. In the experiments, HFIP or the potentiating tetanus was followed by a stimulation of 80, 120, or 200 Hz. The results showed that both HFIP and the tetanus increased power output at high contraction velocities (>75 mm/s) when followed by a train of 80 or 120 Hz (200 Hz resulted in no effects). Mechanical power output was increased maximally by HFIP to 120 and 168% by the tetanus. Furthermore, when HFIP or the tetanus were followed by a train of 80 Hz, the peak power in the power-velocity curve tended to be shifted to a higher velocity.
A single shortening contraction reduces the force capacity of muscle fibers, whereas force capacity is enhanced following lengthening. However, how motor unit recruitment and discharge rate (muscle activation) are adapted to such changes in force capacity during submaximal contractions remains unknown. Additionally, there is limited evidence for force enhancement in larger muscles. We therefore investigated lengthening- and shortening-induced changes in activation of the knee extensors. We hypothesized that when the same submaximal torque had to be generated following shortening, muscle activation had to be increased, whereas a lower activation would suffice to produce the same torque following lengthening. Muscle activation following shortening and lengthening (20 degrees at 10 degrees /s) was determined using rectified surface electromyography (rsEMG) in a 1st session (at 10% and 50% maximal voluntary contraction (MVC)) and additionally with EMG of 42 vastus lateralis motor units recorded in a 2nd session (at 4%-47%MVC). rsEMG and motor unit discharge rates following shortening and lengthening were normalized to isometric reference contractions. As expected, normalized rsEMG (1.15 +/- 0.19) and discharge rate (1.11 +/- 0.09) were higher following shortening (p < 0.05). Following lengthening, normalized rsEMG (0.91 +/- 0.10) was, as expected, lower than 1.0 (p < 0.05), but normalized discharge rate (0.99 +/- 0.08) was not (p > 0.05). Thus, muscle activation was increased to compensate for a reduced force capacity following shortening by increasing the discharge rate of the active motor units (rate coding). In contrast, following lengthening, rsEMG decreased while the discharge rates of active motor units remained similar, suggesting that derecruitment of units might have occurred.
We investigated the effects of low frequency fatigue (LFF) on post-exercise changes in rectified surface EMG (rsEMG) and single motor unit EMG (smuEMG) in vastus lateralis muscle (n = 9). On two experimental days the knee extensors were fatigued with a 60-s-isometric contraction (exercise) at 50% maximal force capacity (MFC). On the first day post-exercise (15 s, 3, 9, 15, 21 and 27 min) rsEMG and electrically-induced (surface stimulation) forces were investigated. SmuEMG was obtained on day two. During short ramp and hold (5 s) contractions at 50% MFC, motor unit discharges of the same units were followed over time. Post-exercise MFC and tetanic force (100 Hz stimulation) recovered to about 90% of the pre-exercise values, but recovery with 20 Hz stimulation was less complete: the 20-100 Hz force ratio (mean +/- SD) decreased from 0.65+/-0.06 (pre-exercise) to 0.56+/-0.04 at 27 min post-exercise (P<0.05), indicative of LFF. At 50% MFC, pre-exercise rsEMG (% pre-exercise maximum) and motor unit discharge rate were 51.1 +/- 12.7% and 14.1 +/- 3.7 (pulses per second; pps) respectively, 15 s post-exercise the respective values were 61.4 +/- 15.4% (P<0.05) and 13.2 +/- 5.6 pps (P>0.05). Thereafter, rsEMG (at 50% MFC) remained stable but motor unit discharge rate significantly increased to 17.7 +/- 3.9 pps 27 min post-exercise. The recruitment threshold decreased (P<0.05) from 27.7 +/- 6.6% MFC before exercise to 25.2 +/- 6.7% 27 min post-exercise. The increase in discharge rate was significantly greater than could be expected from the decrease in recruitment threshold. Thus, post-exercise LFF was compensated by increased motor unit discharge rates which could only partly be accounted for by the small decrease in motor unit recruitment threshold.
The purpose of the present study was to relate the expected inter-subject variability in voluntary drive of the knee extensor muscles during a sustained isometric contraction to the changes in firing rates of single motor units. Voluntary activation, as established with superimposed electrical stimulation was high (range: 91-99%, n=8) during a short maximal contraction, but was lower (range: 69-100%) in most subjects at the point of force failure during a sustained (49.1+/-10.1 s) fatiguing contraction at 50% of maximum force. On a different experimental day the firing behaviour of 27 single motor units was recorded with wire electrodes in the vastus lateralis muscle, 24 of which could be monitored from the time of recruitment to the point of force failure (53.6+/-9.8 s). Motor unit firing behaviour differed considerably among subjects. During the second half of the sustained, fatiguing contraction the changes in firing rate firing rate variability of early recruited units ranged from -10% to +100% and from -50% to +160% respectively among subjects. There were significant positive linear relations between voluntary activation, on the one hand, and rectified surface electromyogram (rsEMG, r=0.82), the changes in motor unit firing rate ( r=0.49) and firing rate variability ( r=0.50) towards the point of force failure on the other. The present data suggest that differences in voluntary drive that appear among subjects during fatigue may be an important determinant of motor unit firing behaviour.
Single motor unit electromyographic (EMG) activity of the knee extensors was investigated at different knee angles with subjects (n = 10) exerting the same absolute submaximal isometric torque at each angle. Measurements were made over a 20 degrees range around the optimum angle for torque production (AngleTmax) and, where feasible, over a wider range (50 degrees ). Forty-six vastus lateralis (VL) motor units were recorded at 20.7 +/- 17.9 %maximum voluntary contraction (%MVC) together with the rectified surface EMG (rsEMG) of the superficial VL muscle. Due to the lower maximal torque capacity at positions more flexed and extended than AngleTmax, single motor unit recruitment thresholds were expected to decrease and discharge rates were expected to increase at angles above and below AngleTmax. Unexpectedly, the recruitment threshold was higher (P < 0.05) at knee angles 10 degrees more extended (43.7 +/- 22.2 N.m) and not different (P > 0.05) at knee angles 10 degrees more flexed (35.2 +/- 17.9 N.m) compared with recruitment threshold at AngleTmax (41.8 +/- 21.4 N.m). Also, unexpectedly the discharge rates were similar (P > 0.05) at the three angles: 11.6 +/- 2.2, 11.6 +/- 2.1, and 12.3 +/- 2.1 Hz. Similar angle independent discharge rates were also found for 12 units (n = 5; 7.4 +/- 5.4 %MVC) studied over the wider (50 degrees ) range, while recruitment threshold only decreased at more flexed angles. In conclusion, the similar recruitment threshold and discharge behavior of VL motor units during submaximal isometric torque production suggests that net motor unit activation did not change very much along the ascending limb of the knee-angle torque relationship. Several factors such as length-dependent twitch potentiation, which may contribute to this unexpected aspect of motor control, are discussed.
When force-velocity-related differences in force capacity were taken into account, neuromuscular activity was not lower during lengthening but was still higher during shortening compared with isometric contractions.
Force and speed parameters were obtained from isometric contractions at different stimulation frequencies of creatine kinase-deficient and wildtype in situ mouse medial gastrocnemius muscles. The absence of creatine kinase did not affect force production at higher stimulation frequencies. However, at frequencies below 140 Hz, forces were lower than the controls (P < 0.05); at the lowest frequency applied (80 Hz) the force was reduced to approximately 60% compared with the wildtype muscles. In contrast, twitch force was not affected. When the contractions were preceded by a brief tetanus (50 ms), the effects of lacking creatine kinase on force production were more pronounced; at 80 Hz stimulation isometric force was further reduced to 66.5 +/- 6.2% (mean +/- SD; n=5) of the single contractions of the deficient muscles and to approximately 42% of the wildtype muscles. Twitch force was now also reduced (by approximately 50%) after the tetanus. The speed of the muscles was not affected in the single contractions. However, after a preceding tetanus, the rate of force rise was reduced by approximately 14% at high frequencies of stimulation. With decreasing frequencies (below 250 Hz), the reduction in speed became more pronounced; at 80 Hz the rate in the creatine kinase-deficient muscles was only 55.2 +/- 3.9% (mean +/- SD; n=5) of the wildtype muscles. No effects of the deficiency were found for the half relaxation times. The data suggest that an impaired creatine kinase system leads to lower activation levels at submaximal stimulation frequencies, possibly by a reduction in Ca2+-release during repetitive stimulation. Similar effects may be expected in normal fatigued muscle when phosphocreatine is depleted.
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