scholarly journals Human stretch reflex pathways reexamined

2014 ◽  
Vol 111 (3) ◽  
pp. 602-612 ◽  
Author(s):  
Ş. Utku Yavuz ◽  
Natalie Mrachacz-Kersting ◽  
Oğuz Sebik ◽  
M. Berna Ünver ◽  
Dario Farina ◽  
...  

Reflex responses of tibialis anterior motor units to stretch stimuli were investigated in human subjects. Three types of stretch stimuli were applied (tap-like, ramp-and-hold, and half-sine stretch). Stimulus-induced responses in single motor units were analyzed using the classical technique, which involved building average surface electromyogram (SEMG) and peristimulus time histograms (PSTH) from the discharge times of motor units and peristimulus frequencygrams (PSF) from the instantaneous discharge rates of single motor units. With the use of SEMG and PSTH, the tap-like stretch stimulus induced five separate reflex responses, on average. With the same single motor unit data, the PSF technique indicated that the tap stimulus induced only three reflex responses. Similar to the finding using the tap-like stretch stimuli, ramp-and-hold stimuli induced several peaks and troughs in the SEMG and PSTH. The PSF analyses displayed genuine increases in discharge rates underlying the peaks but not underlying the troughs. Half-sine stretch stimuli induced a long-lasting excitation followed by a long-lasting silent period in SEMG and PSTH. The increase in the discharge rate, however, lasted for the entire duration of the stimulus and continued during the silent period. The results are discussed in the light of the fact that the discharge rate of a motoneuron has a strong positive linear association with the effective synaptic current it receives and hence represents changes in the membrane potential more directly and accurately than the other indirect measures. This study suggests that the neuronal pathway of the human stretch reflex does not include inhibitory pathways.

1994 ◽  
Vol 76 (6) ◽  
pp. 2411-2419 ◽  
Author(s):  
S. J. Garland ◽  
R. M. Enoka ◽  
L. P. Serrano ◽  
G. A. Robinson

The activity of 50 single motor units was recorded in the biceps brachii muscle of human subjects while they performed submaximal isometric elbow flexion contractions that were sustained to induce fatigue. The purposes of this study were to examine the influence of fatigue on motor unit threshold force and to determine the relationship between the threshold force of recruitment and the initial interimpulse interval on the discharge rates of single motor units during a fatiguing contraction. The discharge rate of most motor units that were active from the beginning of the contraction declined during the fatiguing contraction, whereas the discharge rates of most newly recruited units were either constant or increased slightly. The absolute threshold forces of recruitment and derecruitment decreased, and the variability of interimpulse intervals increased after the fatigue task. The change in motor unit discharge rate during the fatigue task was related to the initial rate, but the direction of the change in discharge rate could not be predicted from the threshold force of recruitment or the variability in the interimpulse intervals. The discharge rate of most motor units declined despite an increase in the excitatory drive to the motoneuron pool during the fatigue task.


1983 ◽  
Vol 49 (2) ◽  
pp. 291-302 ◽  
Author(s):  
B. McKeon ◽  
D. Burke

1. In human subjects, microelectrode recordings were made from 25 muscle spindle afferents and two tendon organ afferents coming from muscles innervated by the peroneal nerve. 2. Stimulation at low intensity through the recording microelectrode activated efferent axons innervating motor units in close proximity to the muscle spindle or tendon organ. There was a clear alteration in the discharge of 17 afferents (15 muscle spindle, 2 tendon organ) in response to twitch contractions that involved only one, two, or three motor units. With three other afferents there was a less overt but statistically significant alteration in discharge rate by the twitch contraction of a single motor unit. 3. The sensitivity of 21 receptors (20 spindles, 1 tendon organ) to twitch contractions of anatomically close motor units was contrasted with their sensitivity to twitches of more remote motor units in the muscle. In no instance was the sensitivity to the contraction of remote motor units greater than that to the contraction of local motor units stimulated through the microelectrode; with remote stimulation many units usually had to be activated before the resulting twitch contraction altered the discharge of an afferent. 4. It is concluded that muscle spindles as well as tendon organs can play a role in monitoring the activity of motor units anatomically close to the receptor.


1985 ◽  
Vol 53 (1) ◽  
pp. 32-42 ◽  
Author(s):  
L. Jami ◽  
J. Petit ◽  
U. Proske ◽  
D. Zytnicki

The discharges of individual tendon organs of peroneus longus and tertius muscles were examined in anesthetized cats during stimulation of single motor units at frequencies that elicit unfused contraction (5-50/s). At these frequencies nearly all the fast-contracting motor units activating a tendon organ elicited responses whose discharge rates reproduced the stimulation frequency ("1:1 driving"), whereas slow-contracting motor units elicited responses in which the discharge rate was higher than the stimulation frequency. When a motor unit stimulated at 40/s developed a gradually potentiating tension, the tendon organ discharge could remain locked on stimulation frequency over an appreciable range of the increasing tension as if the receptor responded to the tension oscillations rather than to the mean level of tension. The only visible effect of the gradual increase in mean tension on the tendon organ response was a gradual decrease of the delay between each stimulus and the corresponding impulse. Driving of tendon organ discharge at the stimulation frequency occurred not only when relatively large oscillations were superimposed on a low level of static tension but also when the static component of the tension was quantitatively preponderant. These observations suggest that during unfused contractions the dynamic component of the stimulus (i.e., oscillation of tension) exerts a prevailing influence on the discharge pattern of tendon organs. Computed simulations of tendon organ responses confirmed that a relatively strong dynamic sensitivity could account for the observed behavior of the receptor.


2014 ◽  
Vol 112 (7) ◽  
pp. 1685-1691 ◽  
Author(s):  
Christopher J. Dakin ◽  
Brian H. Dalton ◽  
Billy L. Luu ◽  
Jean-Sébastien Blouin

Rectification of surface electromyographic (EMG) recordings prior to their correlation with other signals is a widely used form of preprocessing. Recently this practice has come into question, elevating the subject of EMG rectification to a topic of much debate. Proponents for rectifying suggest it accentuates the EMG spike timing information, whereas opponents indicate it is unnecessary and its nonlinear distortion of data is potentially destructive. Here we examine the necessity of rectification on the extraction of muscle responses, but for the first time using a known oscillatory input to the muscle in the form of electrical vestibular stimulation. Participants were exposed to sinusoidal vestibular stimuli while surface and intramuscular EMG were recorded from the left medial gastrocnemius. We compared the unrectified and rectified surface EMG to single motor units to determine which method best identified stimulus-EMG coherence and phase at the single-motor unit level. Surface EMG modulation at the stimulus frequency was obvious in the unrectified surface EMG. However, this modulation was not identified by the fast Fourier transform, and therefore stimulus coherence with the unrectified EMG signal failed to capture this covariance. Both the rectified surface EMG and single motor units displayed significant coherence over the entire stimulus bandwidth (1–20 Hz). Furthermore, the stimulus-phase relationship for the rectified EMG and motor units shared a moderate correlation ( r = 0.56). These data indicate that rectification of surface EMG is a necessary step to extract EMG envelope modulation due to motor unit entrainment to a known stimulus.


1992 ◽  
Vol 67 (5) ◽  
pp. 1375-1384 ◽  
Author(s):  
A. M. Aniss ◽  
S. C. Gandevia ◽  
D. Burke

1. Reflex responses were elicited in muscles that act at the ankle by electrical stimulation of low-threshold afferents from the foot in human subjects who were reclining supine. During steady voluntary contractions, stimulus trains (5 pulses at 300 Hz) were delivered at two intensities to the sural nerve (1.2-4.0 times sensory threshold) or to the posterior tibial nerve (1.1-3.0 times motor threshold for the intrinsic muscles of the foot). Electromyographic (EMG) recordings were made from tibialis anterior (TA), peroneus longus (PL), soleus (SOL), medial gastrocnemius (MG), and lateral gastrocnemius (LG) muscles by the use of intramuscular wire electrodes. 2. As assessed by averages of rectified EMG, stimulation of the sural or posterior tibial nerves at nonpainful levels evoked a complex oscillation with onset latencies as early as 40 ms and lasting up to 200 ms in each muscle. The most common initial responses in TA were a decrease in EMG activity at an onset latency of 54 ms for sural stimuli, and an increase at an onset latency of 49 ms for posterior tibial stimuli. The response of PL to stimulation of the two nerves began with a strong facilitation of 44 ms (sural) and 49 ms (posterior tibial). With SOL, stimulation of both nerves produced early inhibition beginning at 45 and 50 ms, respectively. With both LG and MG, sural stimuli produced an early facilitation at 52-53 ms. However, posterior tibial stimuli produced different initial responses in these two muscles: facilitation in LG at 50 ms and inhibition in MG at 51 ms. 3. Perstimulus time histograms of the discharge of 61 single motor units revealed generally similar reflex responses as in multiunit EMG. However, different reflex components were not equally apparent in the responses of different single motor units: an individual motor unit could respond slightly differently with a change in stimulus intensity or background contraction level. The multiunit EMG record represents a global average that does not necessarily depict the precise pattern of all motor units contributing to the average. 4. When subjects stood erect without support and with eyes closed, reflex patterns were seen only in active muscles, and the patterns were similar to those in the reclining posture. 5. It is concluded that afferents from mechanoreceptors in the sole of the foot have multisynaptic reflex connections with the motoneuron pools innervating the muscles that act at the ankle. When the muscles are active in standing or walking, cutaneous feedback may play a role in modulating motoneuron output and thereby contribute to stabilization of stance and gait.


2008 ◽  
Vol 33 (6) ◽  
pp. 1086-1095 ◽  
Author(s):  
Teatske M. Altenburg ◽  
Cornelis J. de Ruiter ◽  
Peter W.L. Verdijk ◽  
Willem van Mechelen ◽  
Arnold de Haan

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° at 10°/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.


1994 ◽  
Vol 72 (4) ◽  
pp. 1885-1896 ◽  
Author(s):  
E. Smits ◽  
P. K. Rose ◽  
T. Gordon ◽  
F. J. Richmond

1. We depleted single motor units in feline sartorius muscles of glycogen by stimulating their motoneurons intracellularly. We mapped the intramuscular distribution of depleted fibers by inspecting histological cross-sections throughout the length of sartorius. 2. We selected ten depleted motor units for detailed study and quantitative analysis. Nine motor units were located in the anterior head of sartorius. One was located in a muscle whose distal half appeared to have been damaged some time before the acute experiment. A single motor unit was located in the medial head of sartorius. 3. Five motor units were composed of fast-twitch glycolytic (FG) muscle fibers, two of fast-twitch oxidative glycolytic (FOG) muscle fibers, and three of slow-twitch oxidative (SO) muscle fibers. Estimates of the numbers of depleted fibers in motor units of anterior sartorius indicated that FG motor units were larger (mean 566 fibers) than FOG and SO motor units (SO mean 190, FOG mean 156 fibers). The SO motor unit in the damaged muscle had 550 fibers. One motor unit depleted in the medial head of sartorius had 270 fibers with FG profiles. 4. Muscle fibers belonging to each anterior motor unit were never distributed throughout the whole cross-section of anterior sartorius at any proximodistal level. Furthermore, fibers were distributed nonuniformly along the proximodistal axis of the muscle. In most muscles at least a few depleted fibers were found at all proximodistal levels. However, in one normal muscle and the damaged muscle, depleted fibers were confined to the proximal end. 5. The fibers in the medial motor unit were confined to a strip that did not extend across the whole cross-section of the muscle head. Fibers within this strip were scattered quite evenly from origin to insertion. This medial FG motor unit occupied a smaller territory and contained fewer fibers than anterior motor units of the same histochemical type. 6. These results show that sartorius motor units are not distributed uniformly in the mediolateral plane; those in anterior sartorius were distributed asymmetrically in the proximodistal axis as well. This finding has important functional implications for the way in which we model force development and transmission in sartorius and other long muscles.


1977 ◽  
Vol 20 (4) ◽  
pp. 613-630 ◽  
Author(s):  
Harvey M. Sussman ◽  
Peter F. MacNeilage ◽  
Randall K. Powers

Recruitment and discharge patterns of single motor units (MUs) in the anterior belly of digastric were studied during speech in three subjects, using electrodes facilitating selective recording at high force levels. Fixed recruitment order was observed in over 99% of all comparisons. Later recruited units invariably possessed muscle action potentials of higher amplitude, suggesting that units were activated in accordance with the “size principle.” Additional evidence for this was that later recruited units, of a set of three studied during speech, motor unit training, and isometric force ramps, showed greater sensitivity to input, and greater dynamic range than earlier recruited units. Units in this set were much more sensitive to rapid changes in input associated with speech gestures than to static activation even at high force levels. Several significant relations between discharge characteristics and aspects of movement dynamics were observed, including relations between (1) recruitment interval (MU1 to MU3) and latency of mandibular lowering, (2) onset of initial discharge of MU1 and relative mechanical advantage of the mandible, (3) number of MUs active and velocity and displacement of the mandible, and (4) discharge rate of MU3 and velocity and displacement of the mandible.


1990 ◽  
Vol 68 (1) ◽  
pp. 26-34 ◽  
Author(s):  
M. A. Nordstrom ◽  
T. S. Miles

The spike-triggered averaging technique was used to determine the time course and extent of fatigue of single motor unit twitches in the human masseter. This is the first report of a fatigue test having been applied to masseter motor units in either animals or humans. The human masseter was found to be comprised predominantly of fast-twitch motor units with a broad spectrum of fatigability. Very few physiological type S units were found, despite histochemical evidence for a substantial population of type I fibers in the masseter. In addition, there was no significant correlation between fatigability and either twitch amplitude or contractile speed in the motor units studied. The latter observations are consistent with the unusual histological features of the masseter. Comparison with other human fatigue data suggests that the extent of fatigue in the present population of masseter motor units after approximately 3,000 activations is similar to that reported for populations of units in first dorsal interosseous and medial gastrocnemius.


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