Spinal reflexes and coactivation of ankle muscles during a submaximal fatiguing contraction.

This study examined the involvement of spinal mechanisms in the control of coactivation during a sustained contraction of the ankle dorsiflexors at 50% of maximal voluntary contraction. Changes in the surface electromyogram (EMG) of the tibialis anterior and of two antagonist muscles, the soleus and lateral gastrocnemius, were investigated during and after the fatigue task. Concurrently, the compound action potential (M-wave) and the Hoffmann reflex of the soleus and lateral gastrocnemius were recorded. The results showed that the torque of the ankle dorsiflexors and the average EMG of the tibialis anterior during maximal voluntary contraction declined by 40.9 +/- 17.7% (mean +/- SD; P < 0.01) and 37.0 +/- 19.9% (P < 0.01), respectively, at task failure. During the submaximal fatiguing contraction, the average EMG of both the agonist and antagonist muscles increased, leading to a nearly constant ratio at the end of the contraction when normalized to postfatigue values. In contrast to the monotonic increase in average EMG of the antagonist muscles, the excitability of their spinal reflex pathways exhibited a biphasic modulation. The amplitude of the Hoffman reflexes in the soleus and lateral gastrocnemius increased to 147.5 +/- 52.9% (P < 0.05) and 166.7 +/- 74.9% (P < 0.01), respectively, during the first 20% of the contraction and then subsequently declined to 66.3 +/- 44.8 and 74.4 +/- 44.2% of their initial values. In conclusion, the results show that antagonist coactivation did not contribute to task failure. The different changes in voluntary EMG activity and spinal reflex excitability in the antagonist muscles during the fatiguing contraction support the concept that the level of coactivation is controlled by supraspinal rather than spinal mechanisms. The findings indicate, however, that antagonist coactivation cannot simply be mediated by a central descending "common drive" to the motor neuron pools of the agonist-antagonist muscle pairs. Rather, they suggest a more subtle regulation of the drive, possibly through presynaptic mechanisms, to the motoneurons that innervate the antagonist muscles.     

Activation among the elbow flexor muscles differs when maintaining arm position during a fatiguing contraction.

Twenty-four men (n = 11) and women (n = 13) supported an inertial load equivalent to 20% of the maximum voluntary contraction force with the elbow flexor muscles for as long as possible while maintaining a constant elbow angle at 90 degrees. Endurance time did not differ on the three occasions that the task was performed (320 +/- 149 s; P > 0.05), and there was no difference between women (360 +/- 168 s) and men (273 +/- 108 s; P = 0.11). The rate of increase in average electromyogram (EMG) for the elbow flexor muscles was similar across sessions (P > 0.05). However, average EMG during the fatiguing task increased for the long head of biceps brachii, brachioradialis, and brachialis (P < 0.05) but not for the short head of biceps brachii. Furthermore, the average EMG for the brachialis was greater at the start and end of the contraction compared with the other elbow flexor muscles. The rate of bursts in EMG activity increased during the fatiguing contraction and was greater in brachialis (1.0 +/- 0.2 bursts/min) compared with the other elbow flexor muscles (0.5 +/- 0.1 bursts/min). The changes in the standard deviation of acceleration, mean arterial pressure, and heart rate during the fatiguing contractions were similar across sessions. These findings indicate that the EMG activity, which reflects the net excitatory and inhibitory input received by the motoneurons in the spinal cord, was not adaptable over repeat sessions for the maintain-position task. Furthermore, these results contrast those from a previous study (Hunter SK and Enoka RM. J Appl Physiol 94: 108-118, 2003) when the goal of the isometric contraction was to maintain a constant force. These results, from a series of studies on the elbow flexor muscles, indicate that the type of load supported during the fatiguing contraction influences the extent to which endurance time can change with repeat performances of the task.     

Older adults are less steady during submaximal isometric contractions with the knee extensor muscles.

This study compared the steadiness of submaximal contractions with the knee extensor muscles in young and old adults. Twenty young and twenty old subjects underwent assessment of isometric maximum voluntary contraction (MVC), one-repetition maximum (1-RM) strength, and steadiness during isometric, concentric, and eccentric contractions with the knee extensor muscles. The old adults displayed 33% lower MVC force and a 41% lower 1-RM load. The coefficient of variation for force was significantly greater for the old adults during isometric contractions at 2, 5, and 10% of MVC but not at 50% MVC. The decline in steadiness at low forces experienced by the men was marginally greater than that experienced by the women. The steadiness of concentric and eccentric contractions was similar in young and old adults at 5, 10, and 50% of 1-RM load. Old subjects exhibited greater coactivation of an antagonist muscle compared with young subjects during the submaximal isometric and anisometric contractions. These results indicate that, whereas the ability to exert steady submaximal forces with the knee extensor muscles was reduced in old adults, fluctuations in knee joint angle during slow movements were similar for young and old adults.     

Moment distribution among human elbow extensor muscles during isometric and submaximal extension

To understand better how the central nervous system (CNS) distributes a joint moment among muscles, moment distribution among the three heads of the triceps and the anconeus muscles during isometric elbow extension was quantified in vivo and noninvasively. Electrical stimulation was used to activate an individual muscle selectively at various contraction levels, and the relationship between the peak M-wave amplitude and peak elbow extension moment was established across various contraction levels for each muscle. The relationship was then used to calibrate the corresponding EMG signal and determine moment distribution among the muscles during voluntary isometric elbow extension. Results showed that moment distribution among muscles was not proportional to the muscles' physiological cross-sectional areas (PCSA) and the CNS favored uniarticular muscles for the isometric task performed: the uniarticular lateral and medial heads of the triceps were dominant (contributing approximately 70-90% of the total elbow extension moment) and the anconeus contributed significantly, especially at the lower levels of elbow extension moment (up to approximately 15% of the extension moment). In contrast, the two-joint long head of the triceps contributed significantly less than the uniarticular heads of the triceps. While the absolute contributions of all the muscles increased with the total elbow extension moment, the relative contributions of the muscles may increase or decrease with the elbow extension moment. Cross-validation using fresh data (not used in determining the moment distribution) showed close match between the measured and predicted elbow extension moment except for trials in which fatigue became significant.     

Postactivation potentiation, fiber type, and twitch contraction time in human knee extensor muscles.

In small mammals, muscles with shorter twitch contraction times and a predominance of fast-twitch, type II fibers exhibit greater posttetanic twitch force potentiation than muscles with longer twitch contraction times and a predominance of slow-twitch, type I fibers. In humans, the correlation between potentiation and fiber-type distribution has not been found consistently. In the present study, postactivation potentiation (PAP) was induced in the knee extensors of 20 young men by a 10-s maximum voluntary isometric contraction (MVC). Maximal twitch contractions of the knee extensors were evoked before and after the MVC. A negative correlation (r = -0. 73, P < 0.001) was found between PAP and pre-MVC twitch time to peak torque (TPT). The four men with the highest (HPAP, 104 +/- 11%) and lowest (LPAP, 43 +/- 7%) PAP values (P < 0.0001) underwent needle biopsies of vastus lateralis. HPAP had a greater percentage of type II fibers (72 +/- 9 vs. 39 +/- 7%, P < 0.001) and shorter pre-MVC twitch TPT (61 +/- 12 vs. 86 +/- 7 ms, P < 0.05) than LPAP. These data indicate that, similar to the muscles of small mammals, human muscles with shorter twitch contraction times and a higher percentage of type II fibers exhibit greater PAP.     

Differential serial sarcomere number adaptations in knee extensor muscles of rats is contraction type dependent.

Sarcomerogenesis, or the addition of sarcomeres in series within a fiber, has a profound impact on the performance of a muscle by increasing its contractile velocity and power. Sarcomerogenesis may provide a beneficial adaptation to prevent injury when a muscle consistently works at long lengths, accounting for the repeated-bout effect. The association between eccentric exercise, sarcomerogenesis and the repeated-bout effect has been proposed to depend on damage, where regeneration allows sarcomeres to work at shorter lengths for a given muscle-tendon unit length. To gain additional insight into this phenomenon, we measured fiber dynamics directly in the vastus lateralis (VL) muscle of rats during uphill and downhill walking, and we measured serial sarcomere number in the VL and vastus intermedius (VI) after chronic training on either a decline or incline grade. We found that the knee extensor muscles of uphill walking rats undergo repeated active concentric contractions, and therefore they suffer no contraction-induced injury. Conversely, the knee extensor muscles during downhill walking undergo repeated active eccentric contractions. Serial sarcomere numbers change differently for the uphill and downhill exercise groups, and for the VL and VI muscles. Short muscle lengths for uphill concentric-biased contractions result in a loss of serial sarcomeres, and long muscle lengths for downhill eccentric-biased contractions result in a gain of serial sarcomeres.     

Local blood circulation among knee extensor synergists in relation to alternate muscle activity during low-level sustained contraction.

The relation between local circulation and alternate muscle activity among knee extensor synergists was determined during low-level sustained knee extension at 2.5% of maximal voluntary contraction for 60 min in seven subjects. Blood volume of rectus femoris (RF) and vastus lateralis (VL) was assessed by using near-infrared spectroscopy. Surface electromyogram (EMG) was recorded from RF, VL, and vastus medialis (VM). Alternate muscle activity was observed between RF and either VL or VM. Cross-correlation analysis was used to investigate the relation between blood volume and integrated EMG (iEMG) sequences throughout the task. One negative peak in the cross-correlation function was seen between the iEMG and blood volume with time lag of 30-60 s, indicating that muscle activity increases (or decreases) with the decrease (or increase) in local circulation with the corresponding time lag. Two cases in the emergence of alternate muscle activities, i.e., an increase in the EMG of RF accompanied by a decline of EMG in VL (case I) and vice versa (case II) were further analyzed. The time lag between iEMG and blood volume was longer in case I than that in case II. These results were statistically significant in the RF but not in the VL. It is concluded that even during low-level sustained contraction, local circulation is modulated by the alternate muscle activity of knee extensor synergists, and a negative correlation between the muscle activity and blood volume sequences was found in only RF but not in VL.     

Effect of aging on fatigue characteristics of elbow flexor muscles during sustained submaximal contraction.

The purpose of this study was to compare fatigue-related measures of central and peripheral mechanisms between young and elderly subjects for a task performed with elbow flexor muscles. Ten young and nine elderly subjects performed a sustained submaximal fatigue task at 35% of their maximum voluntary contraction torque. Measures of neuromuscular function, reflecting changes in neuromuscular propagation, voluntary activation, excitation-contraction-relaxation processes, and metabolite buildup, were taken before, during, and after the fatigue task. The main results were the absence of neuromuscular propagation failure in either young or elderly subjects, the presence of central fatigue at the end of the fatigue task in 7 of 9 elderly but only 3 of 10 young subjects, and lesser changes in twitch torque contraction-relaxation variables and electromyographic median frequency in elderly compared with young subjects. The lesser fatigue-related changes in twitch contraction speed and median frequency in elderly compared with young subjects could reflect the increase in type I-to-type II fiber area reported with old age. The presence of significant central fatigue can apparently minimize some of the potential differences present in peripheral fatigue sites.     

Force-velocity relations and fiber composition in human knee extensor muscles.

Standardized measurements of dynamic strength of the kneee extensor muscles were performed in 25 healthy male subjects (17-37 yr) by means of isokinetic contractions, i.e., knee extensions with constant angular velocities. Overall variation between double determinations of maximal torque throughout the 90 degrees arc of motion (0 degrees = fully extended leg) averaged 10% for the different constant velocities chosen. At any given angle of the knee the torque produced was higher for isometric than for dynamic contractions. Dynamic torque decreased gradually with increased speed of shortening. Peak dynamic torque was reached at knee angles in the range: 55-66 degrees, with a displacement toward smaller knee angles with higher angular velocities. Correlations were demonstrated between peak torque produced at the highest speed of muscle shortening and percent as well as relative area of fast twitch fibers in the contracting muscle. In addition muscles with a high percentage of fast twitch fibers had the highest maximal contraction speeds. These observations on intact human skeletal muscle are consistent with earlier findings in animal skeletal muscle preparations.     

Total power output generated during dynamic knee extensor exercise at different contraction frequencies.

A novel approach has been developed for the quantification of total mechanical power output produced by an isolated, well-defined muscle group during dynamic exercise in humans at different contraction frequencies. The calculation of total power output comprises the external power delivered to the ergometer (i.e., the external power output setting of the ergometer) and the "internal" power generated to overcome inertial and gravitational forces related to movement of the lower limb. Total power output was determined at contraction frequencies of 60 and 100 rpm. At 60 rpm, the internal power was 18+/- 1 W (range: 16-19 W) at external power outputs that ranged between 0 and 50 W. This was less (P<0.05) than the internal power of 33+/-2 W (27-38 W) at 100 rpm at 0-50 W. Moreover, at 100 rpm, internal power was lower (P<0.05) at the higher external power outputs. Pulmonary oxygen uptake was observed to be greater (P<0.05) at 100 than at 60 rpm at comparable total power outputs, suggesting that mechanical efficiency is lower at 100 rpm. Thus a method was developed that allowed accurate determination of the total power output during exercise generated by an isolated muscle group at different contraction frequencies.     

Respiration during isometric contraction of the flexor or extensor muscles of the arm in humans

An increase in the instantaneous pulmonary flow and a decrease in the respiratory frequency has been found in man as a consequence of isometric contraction of either flexor or extensor muscles of the elbow. The delay of 200 to 900msec between the onset of the muscle contraction and the beginning of the flow increment suggests that the respiratory system increases its activity due to the proprioreceptor activation. As there is no significant difference in the results obtained during flexor or extensor contraction, the contribution of the two different receptor groups appears to be the same.     

Time to task failure and muscle activation vary with load type for a submaximal fatiguing contraction with the lower leg.

The purpose was to compare the time to failure and muscle activation patterns for a sustained isometric submaximal contraction with the dorsiflexor muscles when the foot was restrained to a force transducer (force task) compared with supporting an equivalent inertial load and unrestrained (position task). Fifteen men and women (mean+/-SD; 21.1+/-1.4 yr) performed the force and position tasks at 20% maximal voluntary contraction force until task failure. Maximal voluntary contraction force performed before the force and position tasks was similar (333+/-71 vs. 334+/-65 N), but the time to task failure was briefer for the position task (10.0+/-6.2 vs. 21.3+/-17.8 min, P<0.05). The rate of increase in agonist root-mean-square electromyogram (EMG), EMG bursting activity, rating of perceived exertion, fluctuations in motor output, mean arterial pressure, and heart rate during the fatiguing contraction was greater for the position task. EMG activity of the vastus lateralis (lower leg stabilizer) and medial gastrocnemius (antagonist) increased more rapidly during the position task, but coactivation ratios (agonist vs. antagonist) were similar during the two tasks. Thus the difference in time to failure for the two tasks with the dorsiflexor muscles involved a greater level of neural activity and rate of motor unit recruitment during the position task, but did not involve a difference in coactivation. These findings have implications for rehabilitation and ergonomics in minimizing fatigue during prolonged activation of the dorsiflexor muscles.     

Twitch potentiation is greater after a fatiguing submaximal isometric contraction performed at short vs. long quadriceps muscle length.

Endurance time of a submaximal sustained contraction is longer when the muscle is fatigued in a shortened position. The aim of the present study was to compare central and peripheral mechanisms of fatigue after an isometric contraction of the knee extensor muscles performed at 20% maximal voluntary contraction (MVC) at two knee angles (35 degrees , short length vs. 75 degrees , long length; 0 degrees = full extension) until exhaustion. Eleven men (24 +/- 4 yr) attended two experimental randomized sessions. Endurance time was greater at 35 degrees compared with 75 degrees (974 +/- 457 vs. 398 +/- 144 s; P < 0.001) despite a similar reduction in knee extensor MVC (-28.4 +/- 16.0%, P < 0.001 vs. -27.6 +/- 18.8%, P < 0.001, respectively). Voluntary activation level was similarly depressed after the fatiguing contraction performed at the two muscle lengths (-19 +/- 16.7% at 35 degrees , P < 0.01 vs. -13.7 +/- 14.5% at 75 degrees , P < 0.01). After the fatiguing contraction, peak twitch potentiation was observed only at the short length (+31.8 +/- 17.6% at 35 degrees , P < 0.01 vs. +6.4 +/- 21.3% at 75 degrees , P > 0.05), whereas M-wave properties were similarly altered for the two angles. These results suggest that 1) central fatigue at task failure for a sustained isometric contraction was not dependent on the muscle length, and 2) the longer endurance time of a sustained isometric contraction performed at a shortened length is related to potentiation. It is suggested that the greater endurance time of a sustained isometric contraction observed at 35 degrees is related to the occurrence of potentiation at this short length, because central fatigue is similar at task failure for both tasks.     

Mechanical and morphological properties of the triceps surae muscle-tendon unit in old and young adults and their interaction with a submaximal fatiguing contraction.

The purposes of this study were to examine (a) whether the morphological properties of the muscle gastrocnemius medialis (GM) contribute to the known enhanced muscle fatigue resistance during submaximal sustained isometric plantar flexion contraction of old compared to young adults and (b) whether a submaximal fatiguing contraction differently affects the mechanical properties of the GM tendon and aponeurosis of old and young adults. Fourteen old and 12 young male subjects performed maximal voluntary isometric plantar flexions (MVC) on a dynamometer before and after a submaximal fatiguing task (40% MVC). Moments and EMG signals from the gastrocnemius medialis and lateralis, soleus and tibialis anterior muscles were measured. The elongation of the GM tendon and aponeurosis and the morphological properties of its contractile element were examined by means of ultrasonography. The old adults showed lower maximal ankle joint moment, stiffness and fascicle length in both tested conditions. The submaximal fatiguing contraction did not affect the force-strain relationship of the GM tendon and aponeurosis of either young or old adults. The time to task failure was longer for the old adults and was strongly correlated with the fascicle length (r(2)=0.50, P<0.001). This provides evidence on that the lower ratio of the active muscle volume to muscle force for the old adults might be an additional mechanism contributing to the known age related increase in muscle fatigue resistance.     

Age-related fatigability in knee extensors and knee flexors during dynamic fatiguing contractions

This study investigated the effects of dynamic knee extension and flexion fatiguing task on torque and neuromuscular responses in young and older individuals. Eighteen young (8 males; 25.1 ± 3.2 years) and 17 older (8 males; 69.7 ± 3.7 years) volunteered. Following a maximal voluntary isometric contraction test, participants performed a fatiguing task involving 22 maximal isokinetic (concentric) knee extension and flexion contractions at 60°/s, while surface EMG was recorded simultaneously from the knee extensors (KE) and flexors (KF). Fatigue-induced relative torque reductions were similar between age groups for KE (peak torque decrease: 25.15% vs 26.81%); however, KF torque was less affected in older individuals (young vs older peak torque decrease: 27.6% vs 11.5%; p < 0.001) and this was associated with greater increase in hamstring EMG amplitude (p < 0.001) and hamstrings/quadriceps peak torque ratio (p < 0.01). Furthermore, KE was more fatigable than KF only among older individuals (peak torque decrease: 26.8% vs 11.5%; p < 0.001). These findings showed that the age-related fatigue induced by a dynamic task was greater for the KE, with greater age-related decline in KE compared to KF.     

Force fluctuations are modulated by alternate muscle activity of knee extensor synergists during low-level sustained contraction.

The study examined the hypothesis that altered synergistic activation of the knee extensors leads to cyclic modulation of the force fluctuations. To test this hypothesis, the force fluctuations were investigated during sustained knee extension at 2.5% of maximal voluntary contraction force for 60 min in 11 men. Surface electromyograms (EMG) were recorded from the rectus femoris (RF), vastus lateralis (VL), and vastus medialis (VM) muscles. The SD of force and average EMG (AEMG) of each muscle were calculated for 30-s periods during alternate muscle activity. Power spectrum of force was calculated for the low- (< or =3 Hz), middle- (4-6 Hz), and high-frequency (8-12 Hz) components. Alternate muscle activity was observed between RF and the set of VL and VM muscles. The SD of force was not constant but variable due to the alternate muscle activity. The SD was significantly greater during high RF activity compared with high VL and VM activity (P < 0.05), and the correlation coefficient between the SD and AEMG was significantly greater in RF [0.736 (SD 0.095), P < 0.05] compared with VL and VM. Large changes were found in the high-frequency component. During high RF activity, the correlation coefficient between the SD and high-frequency component [0.832 (SD 0.087)] was significantly (P < 0.05) greater compared with other frequency components. It is suggested that modulations in knee extension force fluctuations are caused by the unique muscle activity in RF during the alternate muscle activity, which augments the high-frequency component of the fluctuations.