Quantitative analysis of reflex responses in the averaged surface electromyogram.

The relationship between the averaged surface electromyogram (EMG) and the activity of motoneurones in reflex studies is analysed mathematically. This analysis reveals that, subject to certain conditions being met, the integral of the average of the unrectified EMG is linearly related to the activity of motor units in the muscle. This was tested with experimental data. The activity of 2 motor units, and the surface EMG, were recorded directly in a reflex paradigm. A close match was found between the integral of the average of the unrectified EMG and the summed peristimulus time histograms (PSTHs) of the units' activity. This analysis gives more quantitative measurements of the timing of motoneuronal activity in reflexes than the conventional analysis of the EMG. It also offers the potential for making quantitative measurements of changes in motor unit activity evoked by different stimuli. This analysis avoids the pitfall of the artefactual peaks that can occur in the average of the rectified EMG as a consequence of the full-wave rectification. It is concluded that the integral of the unrectified average offers a valuable adjunct to the conventional analysis in many reflex studies. The analysis can also be applied to quantitative studies of neurograms.     

Spinal stretch reflex and cortical evoked potential amplitudes versus muscle stretch amplitude in the monkey arm

While investigating operant conditioning of the primate spinal stretch reflex (SSR), we studied SSR amplitude and cortical somatosensory evoked potential (SEP) amplitude as stretch amplitude changed in the monkey arm. Initial muscle length and background EMG activity remained constant. With change in stretch amplitude (and proportional change in stretch velocity and acceleration), changes in SSR and SEP amplitudes were respectively 0.75 and 0.66 as great. The lesser change in SSR amplitude may reflect saturation of Ia afferents, while that in SEP amplitude may also reflect participation of other peripheral receptors.     

Quantitation of the stretch reflex

The stretch reflex should ideally be quantitated for better clinical use by standardizing the muscle stretch and measuring the resulting muscle contraction. Quantitation of muscle contraction can be done by force measurements or electromyographic recordings. The electromyographic response to stretch consists of one component (short latency response) for short stretches (less than 15 ms) and of 2 or 3 components (short and long latency responses) for longer stretches (greater than 40-50 ms). The magnitude of the phasic stretch reflex is reflected by the short latency response, whereas the magnitude of the tonic stretch reflex is reflected by both the short and the long latency responses. In clinical studies of upper motor neuron syndromes, the knee jerk and the muscle tone correlated with the magnitude of the short latency response. In patients with paralysis agitans an increased long latency response, which correlated to the rigidity, was found.     

Behavior of the stretch reflex in a multi-jointed limb

Local microiontophoretic administration of cholecystokinin octapeptide (CCK) increased the firing rates of neurons in the dorsomedial nucleus accumbens (NAc), but exerted little to no effect on lateral NAc neurons. This regionally defined CCK-effect corresponds to the topographical distribution of CCK-like immunoreactive nerve terminal fiber networks and CCK receptors within the NAc. The excitatory effects of CCK were selectively antagonized by the CCK antagonist proglumide. Dopamine (DA) decreased the firing of NAc cells and reversed CCK-induced excitation. These results suggest that CCK and DA may interact to influence the activity of neurons within the dorsomedial NAc.     

Continuous wavelet transform in the evaluation of stretch reflex responses from surface EMG

OBJECTIVE: This is the first reported use of the continuous wavelet transform (CWT) of the surface EMG (sEMG) to extract the reflex response to muscle stretch. We used a modulus-based method to estimate instantaneous amplitude-envelopes from ridges of the CWT (referred in this work as sEMG intensity) to extract the dynamic reflex response from sEMG. We tested the method on tendon reflexes where excellent temporal resolution is required to identify the different latency components, and on the tonic stretch reflex (tonic SR) response to an ongoing perturbation that characteristically has a low signal to noise ratio. METHODS: Eight subjects without neurological impairment were subjected to a series of archilles tendon taps and a 2 min continuous perturbation of the ankle using a pseudo-sinusoidal stretch profile containing frequencies from 0.1 to 8.0 Hz. The tendon reflexes were assessed in the soleus muscle at 10% of MVC and the tonic SR in tibialis anterior while the muscle was relaxed, at 5 and 10% of maximal voluntary contraction. Root mean square (RMS) and wavelet ridge extraction was applied to the sEMG signal to extract sEMG amplitudes (RMS) and intensities for all reflexes. To obtain the tonic SR, these estimates and those from the sEMG-RMS were subsequently cross-correlated with the perturbation record to yield 2 sets of estimates of reflex gain and coherence for comparison. RESULTS: The sEMG intensities were highly correlated with the torques resulting from a ramped voluntary contraction. Following tendon taps, the method resolved the M1, M2, M3 response components at accurate latencies and with more complete reconstruction of the components than RMS-derived estimates. The wavelet ridge estimates extracted the tonic SR from resting and contracting muscles with significantly higher coherence than RMS estimates. Reflex gain, when estimated from sEMG intensity or sEMG-RMS, demonstrated similar relationships to the perturbation frequency and background contraction level. When the sEMG intensity reflex gain estimates from different subjects were pooled, they showed significantly lower variance about the mean than gain estimates derived from the rectified sEMG. CONCLUSIONS: Wavelet-ridge extraction provides a valid approach to reflex evaluation from sEMG that does not depend on the absolute amplitude of the potentials measured at the EMG electrodes. This may have substantial advantages in more directly comparing responses between subjects on an absolute frequency scale without the need for normalisation against maximal contraction levels.     

Differential effects of plantar cutaneous afferent excitation on soleus stretch and H‐reflex

Previous studies have demonstrated that plantar cutaneous afferents can adjust motoneuron excitability, which may contribute significantly to the control of human posture and locomotion. However, the role of plantar cutaneous afferents in modulating the excitability of stretch and H‐reflex with respect to the location of their excitation remains unclear. In the present study, it was hypothesized that electrical stimulation delivered to the sole of the foot might be followed by modulation of spinal excitability that depends on: (1) the stimulation location and (2) the reflex studied. In these experiments, conditioned and unconditioned stretch and H‐reflexes were evoked in 16 healthy subjects in a seated position. Both reflexes were conditioned by non‐noxious electrical plantar cutaneous afferent stimulation at two different sites, the heel and metatarsal regions, at four different conditioning–test (CT) intervals. The conditioning stimulation delivered to the heel caused a significant facilitation of the soleus stretch reflex for all CT intervals, whereas the soleus H‐reflex had significant facilitation only at CT interval of 50 ms and significant inhibition at longer CT intervals. Stimulation delivered to the metatarsal region, however, resulted mainly in reduced stretch and H‐reflex sizes. This study extends the reported findings on the contribution of plantar cutaneous afferents within spinal interneuron reflex circuits as a function of their location and the reflex studied. Muscle Nerve, 2008     

Diurnal rhythm in the spinal stretch reflex

We studied primate spinal stretch reflex (SSR) amplitude as a function of time of day. SSR amplitude was greatest around midnight and smallest around noon. The diurnal rhythm was not simply a function of number of trials, or of the lighting cycle. This rhythm offers an opportunity to study the neuronal and synaptic mechanisms producing a diurnal change in CNS function. Its existence indicates that the CNS response to a given limb disturbance, and thus the CNS activity underlying a given performance, varies with time of day.     

Adaptive plasticity in the spinal stretch reflex

Monkeys can change the amplitude of the spinal stretch reflex without change in initial α motorneuron tone, as measured by EMG, or in initial muscle length. Change is apparent in 5–10 days, continues to develop over weeks, and persists during inactive periods. Spinal stretch reflex change may be a valuable system for studying the neuronal and synaptic bases of an adaptive change in primate CNS function.     

Central control components of a 'simple' stretch reflex

The monosynaptic stretch reflex is a fundamental feature of sensory-motor organization in most animal groups. In isolation, it serves largely as a negative feedback devoted to postural controls; however, when it is involved in diverse movements, it can be modified by central command circuits. In order to understand the implications of such modifications, a model system has been chosen that has been studied at many different levels: the crayfish walking system. Recent studies have revealed several levels of control and modulation (for example, at the levels of the sensory afferent and the output synapse from the sensory afferent, and via changes in the membrane properties of the postsynaptic neuron) that operate complex and highly adaptive sensory-motor processing. During a given motor task, such mechanisms reshape the sensory message completely, such that the stretch reflex becomes a part of the central motor command.     

Effects of the corticotropin-releasing factor (CRF) on rectal afferent nerves in humans

Abstract Corticotropin-releasing factor (CRF) released in the gastrointestinal mucosa from immune cells or enterochromaffin cells may play a role in the modulation of rectal afferent function. In the current study we evaluated the effects of peripherally administered CRF on afferent mechanisms in the human rectum. We used rectal balloon distention in seven healthy volunteers to evaluate the effect of CRF (1 μ/kg) on visceral afferents originating in the rectum which are involved in the following functions: thresholds and intensity of conscious perception, receptive relaxation, reflex inhibition of internal anal sphincter and a viscerosomatic reflex. Rectal mechanoreceptors were stimulated either by distending the rectum using a volume ramp (40 and 400 mL/min), or by intermittent phasic distention. CRF decreased the thresholds and increased the intensity for the sensation of discomfort in response to both ramp and phasic distention. During slow ramp distention, CRF also lowered the stool threshold. CRF increased rectal compliance during slow ramp distention without affecting the rate of receptive relaxation or the inflection point of the compliance curve. CRF had no effect on viscerosomatic referral patterns, or on the rectoanal inhibitory reflex. These findings are consistent with a dual effect of CRF on afferent pathways mediating perception of aversive rectal sensations, and on rectal smooth muscle.     

Measurements of excitatory postsynaptic potentials in the stretch reflex of normal subjects and spastic patients.

The patellar tendon was tapped by random impulses of triangular waveform and motor unit spikes were recorded from the quadriceps femoris muscle. The cross-correlogram of the taps and the motor unit spikes revealed a primary correlation kernel, the width of which was interpreted as an indicator of the mean time-to-peak of excitatory postsynaptic potentials (EPSPs) elicited monosynaptically in an alpha-motoneurone by the triangular taps. The mean time-to-peak was 7.6 +/- 1.3 ms in normal subjects and 9.0 +/- 1.8 ms in spastic patients (P less than 0.005). The prolonged time-to-peak of EPSP in spastic patients is consistent with the hypothesis that as a result of degeneration of the corticomotoneuronal tract the Ia axons sprout and form more synaptic contacts on distal portions of the dendrites of alpha-motoneurones.     

Phasic stretch reflex of the abdominal muscles

This analysis of the abdominal stretch reflex (ASR) evoked by a tap to the abdomen was designed to explore how abdominal motoneurons process signals from respiratory and nonrespiratory sources. We recorded surface EMGs from the external and internal oblique muscles in standing subjects. Amplitudes of the abdominal stretch reflex varied despite constant tap forces, but strong taps evoked a larger reflex than weak taps. Trunk rotation toward the recording side, or voluntary contraction of the external and internal oblique muscles increased the reflex amplitudes, whereas contralateral rotation reduced their occurrence. An abdominal stretch reflex during a voluntary contraction was followed by a silent period of 40 to 80 ms. Often a late wave followed a reflex by 20 to 40 ms. Amplitudes during breathholds at residual lung volume were larger than those evoked during a breathhold at functional residual capacity, suggesting that abdominal stretch reflex amplitudes are inversely proportional to static lung volume. During quiet breathing the reflex amplitude reached a maximum slightly before end-expiration and decreased progressively to a minimum close to end-inspiration. During rebreathing, background abdominal activity was augmented with highest activity in late expiration. Abdominal stretch reflex amplitudes continued to wax and wane in phase with respiration, and the maximal reflex occurred progressively earlier in expiration. In summary, the abdominal stretch reflex reflects strong control from abdominal muscle spindles, lung proprioceptors, and chemoreceptors. The relative contributions of these inputs need to be determined.     

Stretch reflex variation in the relaxed and the pre-activated quadriceps muscle of normal humans

The stretch reflex to patellar tendon taps was quantified by force measurements at the ankle in 7 normal subjects. In each experiment the stretch reflex was elicited from 14 consecutive stretches by two types of hammers (an ordinary hand-held hammer and a motorized hammer) in the relaxed and the pre-activated quadriceps muscle. The coefficient of variation for the 14 stretch reflexes fell from 54% in the relaxed muscle to 39% in the contracting muscle (p less than 0.05). The stretch reflex varied less between the different levels of active contraction than between the relaxed state and the contracted level (p less than 0.05). The experiment was repeated in all subjects and at all contraction levels. The coefficient of variation of the mean for the two experiments fell non-significantly from a mean value of 27% in the relaxed muscle to 14% in the contracting muscle. The coefficients of variation for the two hammers were almost identical. It is concluded that quantification of the stretch reflex can be made more precisely in the contracting muscle.     

Interaction between voluntary contraction and tonic stretch reflex transmission in normal and spastic patients

Transmission characteristics of tonic stretch reflex (TSR) pathways in both normal and spastic patients have been measured during different levels of sustained voluntary contraction of the biceps brachii muscle. A cross-correlation technique of analysis was used to separate reflex responses from the total electromyographic activity. TSR transmission in normal subjects was observed to change with the level of voluntary contraction; sensitivity to stretch increased approximately three-fold as the subject stiffened the arm by simultaneously contracting flexor and extensor muscles. In contrast, TSR transmission did not alter during voluntary contraction in spastic patients. It is proposed that in spastic patients supraspinal modulation of reflex transmission is impaired because hypersensitive spinal reflexes short-circuit long loop pathways.     

The afferent origin of the secondary somatosensory evoked potential from the lower limb in humans

The afferent origin of the secondary somatosensory evoked potential elicited from stimulation of the sural and tibial nerves was investigated as the limb was cooled. It was hypothesized that the peak of this potential is initiated from primary afferents in the A alpha group. We conclude that the peak of the secondary SEP arises from an afferent source whose diameter is of similar size to that of large diameter A alpha afferents.