Older adults can maximally activate the biceps brachii muscle by voluntary command.

Because some of the decline in strength with age may be explained by an impairment of muscle activation, the purpose of this study was to determine the activation level achieved in biceps brachii by older adults during a maximum voluntary contraction (MVC). This capability was assessed with two superimposition techniques: one calculated the activation level that was achieved during an MVC, and the other provided an estimate of the expected MVC force based on extrapolation with submaximal forces. The activation level in biceps brachii was incomplete (< 100%) for the young (n = 16) and elderly (n = 16) subjects, with the elderly subjects exhibiting the greater deficit. In contrast, there was no difference between the measured and expected MVC forces for either group of subjects, whether the extrapolation involved a third-order polynomial or linearization of the data. Because of the lower signal-to-noise ratio associated with the measurement of activation level and the greater number of measurements that contributed to the estimate of the expected MVC force, we conclude that the older adults were able to achieve complete activation of the biceps brachii muscle during an MVC.

Effect of pronation and supination tasks on elbow flexor muscles.

The aim of this study was to verify the presence of modulatory effects of pronation (P) and supination (S) on biceps brachii (BB) and brachioradialis (BR) electromyographic (EMG) signals while performing isometric elbow flexions at different angles (50, 90, and 130°). The EMG activity of BB and BR was recorded for normal subjects (N = 6) with surface electrodes during an isometric ramp contraction of elbow flexion (F) that was combined with 20% of maximal voluntary contraction (MVC) in pronation or in supination. The results indicate that (a) the BB muscle presents an increased EMG activity in the combined task of S-F and a decreased EMG activity in the combined task of P-F (Friedman ANOVA, p < 0.01); (b) the EMG activity of the BR is not significantly influenced by the different tasks (Friedman's ANOVA, p > 0.05); and (c) the modulation of the BB occurs only at a midrange angle of flexion. This study partly supports results that were previously reported by other authors, suggesting that a combined task can modulate the EMG activity of elbow flexor muscles. However, this modulation, observed especially for the BB, appears to be limited to elbow flexion angles that are close to 90°.

Cocontraction of the elbow muscles during combined tasks of pronation-flexion and supination-flexion.

The aim of this study was to determine if the antagonist activity of the triceps brachii (TB) and anconeus (AN) muscles is modulated when the activity of the biceps brachii (BB) and brachioradialis (BR) is modulated by the performance of combined tasks and to verify if this behavior is similar at different elbow angles. Electromyographic (EMG) activity of BB, BR, AN and TB was recorded for normal subjects (N = 6) with surface electrodes during a ramp isometric contraction in elbow flexion (F) which was performed alone or combined with 20% of maximal voluntary contraction (MVC) in pronation (P) or in supination (S). Two cocontraction ratios, using the EMG root mean square (rms) values of each muscle and identified as BB/TB and BR/AN were calculated. The results indicate that for low flexion torque levels, the BB/TB ratio is higher for the S-F condition while the BR/AN ratio is higher during the pure flexion task. Variations of the EMG activity across tasks were significant only for BB (Friedman ANOVA, p less than .01) whereas there was no significant change in EMG activity (rms) for TB, BR and AN (Friedman ANOVA, p greater than .01). Furthermore, the behavior of both ratios across tasks was similar at 50 degrees, 90 degrees and 130 degrees of elbow flexion. Thus, for isometric conditions, there appears to be no evidence of modulation of EMG activity of elbow extensors while performing combined tasks of S-F and P-F. In addition, cocontraction activity during these tasks tends to be similar across elbow angles.

Wrist muscle activation patterns and stiffness associated with stable and unstable mechanical loads.

The objectives of this study were to examine the effects of load mechanical characteristics and agonist-antagonist muscle cocontraction, on the stretch reflex response of wrist flexor muscles, and to measure the associated wrist stiffness. Subjects were required to maintain a constant wrist angle while operating against flexor loads with different stability characteristics (constant, elastic or unstable). We measured the stretch reflex responses and joint stiffness by applying step displacements of 3 degrees and 10 degrees. Subjects used very little cocontraction of wrist flexor and extensor muscles when the load was constant or elastic, but increased cocontraction dramatically when the load was unstable, in order to increase the wrist stiffness. Although the magnitude of stretch reflex responses also increased with cocontraction, this simply reflected the level of tonic flexor muscle activity. We found no evidence to suggest that phasic stretch reflexes contributed significantly to the joint stiffness in this task. Clear differences in flexor muscle synergy were observed in the presence and absence of cocontraction, particularly when comparing the FCR and FCU muscles.

Regulation of soleus muscle spindle sensitivity in decerebrate and spinal cats during postural and locomotor activities.

1. In order to study fusimotor control in reduced preparations, soleus muscle spindle afferents were recorded in premammillary decerebrate cats (n = 15) during crossed extensor reflexes and, after spinalization, during locomotion produced by either clonidine or L-beta-3,4-dihydroxyphenylalanine (L-DOPA). The soleus muscle was oscillated sinusoidally (0.25 mm, 4 Hz) and the afferent mean firing rate and modulation were calculated. An increase in firing rate was assumed to arise from activity in dynamic gamma-motoneurones (dynamic gamma-drive) when associated with an increase in modulation to stretching, and in static gamma-motoneurones (static gamma-drive) when modulation decreased. 2. At rest in all preparations the firing rate and modulation in primary muscle spindle afferents were generally much higher than after de-efferentation (ventral root section), suggesting a predominant dynamic gamma-drive. Clonidine decreased and even eliminated this presumed resting gamma-drive in many afferents, both in the decerebrate (7 of 8) and the spinal (6 of 18) state. This effect on gamma-drive may account, at least in part, for its suppressive effect on spasticity in humans. 3. When locomotion commenced in clonidine-treated spinal cats, primary afferents generally fired with much higher mean rates (+121%) and lower sensitivities (-32%), suggesting a large increase in static gamma-drive (possibly accompanied by a small decrease in dynamic gamma-drive). These high rates were usually maintained tonically throughout the step cycle. However, a third of the afferents were silenced during locomotor contractions, and de-efferentation had no significant effect on their firing rates. Thus, for some spindles alpha-activity can occur without significant gamma-drive. 4. During locomotion in L-DOPA-treated spinal cats the inferred static gamma-drive only occurred phasically, coactivated with the EMG, though it could precede the EMG by 100-500 ms. In the flexion phase both the afferent rate and modulation were lower than before locomotion, suggesting a lack of effective gamma-drive. 5. Crossed extensor reflexes in decerebrate cats also produced a substantial increase in primary afferent firing rate (+187%) and decrease in sensitivity (-37%), again suggesting increased static gamma-drive (n = 18). This gamma-drive was largely independent of EMG activity and often occurred without alpha-activity. The mean firing rate of secondary muscle spindle afferents increased significantly during locomotion (with L-DOPA) and crossed extensor reflexes, again indicating increased static gamma-drive. Clonidine reduced or eliminated the gamma-drive in seven of eight afferents during crossed extensor reflexes. 6. In conclusion, although there are some common features, such as a predominant static gamma-drive in all walking preparations, the pattern of static and dynamic gamma-drive is not closely linked to alpha-activity under the conditions studied. As well as gamma-drive without alpha-activity, we have shown for the first time that alpha-motoneurones can be activated without significant gamma-drive to many spindles during behavioural tasks.

Mechanism for reflex reversal during walking in human tibialis anterior muscle revealed by single motor unit recording.

1. A reversal in the sign of a cutaneous reflex during walking was recently described in the human. Such reversals were most clearly seen in muscles that were active in two parts of the step cycle, such as the tibialis anterior (TA). The current study determined whether the reversal resulted from differential activation of a single group of motor units. 2. Single motor units were recorded from the TA muscle of healthy human subjects while they walked on a treadmill with a splint that limited motion of the ankle joint. The majority of motor units from which recordings were made (43 out of 46) were active in both the swing phase and the transition from swing to stance, indicating that the two bursts of activity from the TA muscle do not represent the activity of two separate populations of motor units. 3. The firing behavior of three motor units was observed during walking steps when stimuli were applied to the posterior tibial nerve during either the swing phase or the transition from swing to stance. The post-stimulus time histograms indicated that the same motor unit was excited during the swing phase, and inhibited during the transition from swing to stance. 4. The results support the hypothesis that there are parallel excitatory and inhibitory pathways from cutaneous afferents to single motoneurones of the TA muscle. A shift in balance between the two pathways as a function of the step cycle most probably generates the reflex reversal observed.

Gain of the triceps surae stretch reflex in decerebrate and spinal cats during postural and locomotor activities.

1. The triceps surae (TS) stretch reflex was measured in decerebrate cats during crossed extensor stimulation and after spinalization during rhythmic locomotor activity induced by clonidine and manual perineal stimulation. The TS force in response to sinusoidal stretch was measured at a given contraction level before and after deafferentation, and the 'reflex force' was computed by subtracting these two responses. Reflex 'gain' was computed as the ratio of the reflex and deafferented force responses (a unitless estimate of the open loop feedback gain). 2. Prior to locomotion the spontaneous muscle activity was low (less than 15% of maximum), but the reflex gain was relatively high (close to 1.0 with a 5 Hz stretch). When locomotion commenced the reflex gain was markedly lowered when measured at the same contraction level as before locomotion (25% of the gain prior to locomotion). At higher contraction levels the reflex gain was not significantly increased. The reflex force and EMG responses to stretch increased with the contraction level, but their effect on the total reflex gain was cancelled by an associated increase in the intrinsic muscle stiffness. 3. In the decerebrate cat, during weak tonic contractions (spontaneous), the reflex gain was high and comparable with the gain in the resting spinal cat. However, with increased tonic contractions produced by crossed extensor stimulation the reflex gain dropped. At higher contraction levels the gain was not significantly different from the gain during spinal locomotion. 4. When the frequency of stretch was increased from 3 to 20 Hz, EMG responses to stretch increased, but the reflex force decreased, since a more fused contraction developed with the more frequent reflex activations. Overall, the reflex gain decreased with frequency in both spinal and decerebrate cats. The phase lag of the reflex force, relative to the intrinsic muscle force, increased with increasing frequency, due to reflex delays, with a 180 deg lag occurring between 12 and 18 Hz (tremor frequencies). The mean gain was significantly lower and the phase lag was significantly greater during locomotion than during tonic crossed extensor contractions, suggesting different reflex mechanisms. 5. In conclusion, during locomotion in spinal cats afferent feedback from low frequency ankle movements, similar to those occurring during the normal step cycle, reflexly produces a small but significant fraction of the extensor force (about a quarter of the stretch-related force modulation). This fraction is remarkably constant at the different contraction levels of the step cycle. Afferent feedback during higher frequency movement is less effective, minimizing the chance of instability and tremor. In contrast during tonic contractions afferent feedback produces half of the total muscle force during perturbations, clearly contributing to the maintenance of posture.

Stretch and H reflexes in triceps surae are similar during tonic and rhythmic contractions in high decerebrate cats.

During locomotion in decerebrate and spinal cats the group Ia afferents from hind leg muscles are depolarized rhythmically. An earlier study concluded that this locomotor-related primary afferent depolarization (PAD) does not contribute to modulation of monosynaptic reflex pathways during locomotion. This finding indicated that the neural network generating the locomotor rhythm, the central pattern generator (CPG), does not presynaptically inhibit monosynaptic reflexes. In this investigation we tested this prediction in decerebrate cats by measuring the magnitude of reflexes evoked in ankle extensor muscles during periods of tonic contractions and during sequences of rhythmic contractions. The latter occurred when the animal was induced to walk on a treadmill. At the similar levels of activity in the soleus muscle there was no significant difference in the magnitude of the soleus H reflex in these two behavioral situations. Similar results were obtained for reflexes evoked by brief stretches of the soleus muscle. We also examined the reflexes evoked by ramp-and-hold stretches during periods of rhythmic and tonic activity of the isolated medial gastrocnemius (MG) muscle. At similar levels of background activity, the reflexes evoked in the MG muscle were the same during rhythmic and tonic contractions. Our failure to observe a reduction in the magnitude of H reflexes and stretch reflexes during rhythmic contractions, compared with reflexes evoked at the same level of background activity during tonic contractions, is consistent with the notion that the CPG for stepping does not presynaptically inhibit monosynaptic reflexes during the extension phase of locomotor activity. Our results indicate that presynaptic inhibition of the monosynaptic reflex associated with normal locomotion in cats or humans arises from sources other than the extensor burst generating system of the central pattern generator.

Effects of TENS and topical skin anesthesia on soleus H-reflex and the concomitant influence of skin/muscle temperature.

The purpose of this study was to determine, in ten healthy subjects, the extent of soleus motoneuronal excitability during conditions of increased (transcutaneous electrical nerve stimulation [TENS]), decreased (Xylocaine [lidocaine]a anaesthesia) and normal (placebo anaesthesia) cutaneous inputs. Increased cutaneous activity was evoked using a TENS unit, with the two pairs of electrodes placed respectively over the Achilles (S2 dermatome) and tibialis anterior (L5 dermatome) tendons. Experimental and placebo topical anaesthesia were obtained after rubbing Xylocaine (5%) and Vaselineb ointment, respectively, on the skin surface overlying the Achilles tendon. Sets of ten H-responses (Hmax/2) were evoked at a frequency of 1 shock/30s and averaged at regular time intervals before, during and after the testing conditions. The results showed a gradual increase (up to 40% after 20 minutes) of H-reflex amplitude during TENS regardless of whether it was applied on the L5 or S2 dermatome. Furthermore, placebo anesthesia (Vaseline) caused the same gradual facilitatory response (up to 100% after 50 minutes) as that obtained during Xylocaine anaesthesia. Power spectral analysis of the H-responses obtained over time showed that the increase in the peak-to-peak H-response value was accompanied by a shift of the spectral content toward low frequencies. This shift occurred concomitantly with a cooling of the skin overlying the soleus muscle.