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Abnormal development of biceps brachii phasic stretch reflex and persistence of short latency heteronymous reflexes from biceps to triceps brachii in spastic cerebral palsy.

Co-contraction of antagonist muscles is characteristic of spasticity arising from perinatal brain damage but not in spasticity occurring after brain damage in adulthood. Such co-contraction is a normal feature of early post-natal motor development. Heteronymous, monosynaptic Group Ia projections from biceps brachii to both the antagonist triceps brachii and to other synergist and non-synergist muscles of the upper limb occur in the newborn baby and become restricted during the first 4 years to motor neurons of primarily synergistic muscles. Longitudinal and cross-sectional studies have been performed to test the hypothesis that inappropriate heteronymous excitatory projections persist in children with perinatal brain damage who develop spasticity. Subjects with spasticity, from brain damage acquired in adulthood were also studied to determine if these projections simply become unmasked as part of spasticity, independent of the age of occurrence of the brain damage. Twenty-nine healthy newborn babies and 29 at high risk for cerebral palsy, 12 of whom developed spastic quadriparesis, were studied longitudinally for 4 years. Thirty-eight subjects, aged 8-30 years, with spasticity of perinatal origin (11 hemiplegic, 11 quadriplegic, 16 with Rett syndrome) and 11 subjects with stroke in adulthood and spastic hemiplegia were also studied. The results were compared with those obtained in 372 normal subjects aged from birth to 55 years. Small taps were delivered to the tendon of biceps brachii using an electromechanical tapper. Surface EMG was recorded from biceps and triceps brachii, pectoralis major and deltoid. In the longitudinal study, those developing spastic quadriparesis showed persistent low thresholds for the homonymous phasic stretch reflex, which had abnormally short onset latencies. There was persistence of short onset heteronymous excitatory responses in triceps brachii, while a normal pattern of restriction of heteronymous responses to pectoralis major and deltoid occurred. The same pattern was observed in older subject groups with spasticity of perinatal origin. In adults with hemiplegia following stroke the threshold of the homonymous phasic stretch reflex was low, but it had a normal onset latency. There was no evidence of abnormal heteronymous excitatory responses. In conclusion, exaggerated excitatory responses to primary muscle afferent input were observed in the homonymous (biceps brachii) and antagonist (triceps brachii) motor neurons in subjects with spasticity arising from perinatal brain damage. They are likely to play an important role in the predominant co-contraction of agonist/antagonist muscles during voluntary movement observed in subjects with spastic cerebral palsy.

Adolescent↗

Reinnervated muscles fail to produce stretch reflexes.

1. We studied the stretch-evoked reflex organization of hind limb muscles in two decerebrate cats 36 mo after unilateral section and immediate surgical repair of the common nerve supplying the lateral gastrocnemius (LG) and soleus (S) muscles. 2. The production of considerable reflex force by reinnervated muscles in response to electrical stimulation of uninjured nerves indicated substantial functional recovery of motor units. However, reduction in the responsiveness of reinnervated muscles to stretch of the untreated medial gastrocnemius (MG) muscle indicated some deficit in recovery of normal synaptic integration. 3. Stretch failed to elicit autogenic excitation of the reinnervated S and LG. This failure was observed whether the reinnervated muscles were quiescent or contracting in other reflexes. 4. The heterogenic reflex organization of reinnervated muscles was abnormal. Stretch of the reinnervated S failed to evoke heterogenic reflexes both in the untreated MG and in the reinnervated LG. Stretch of the reinnervated LG failed to produce excitation of MG. 5. These findings demonstrate deficiencies in proprioceptive feedback from reinnervated muscles and lead us to expect incomplete recovery of motor function after nerve section.

Afferent Pathways↗

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.

Electromyography↗

Comparison of stretch reflexes and shortening reactions in activated normal subjects with those in Parkinson's disease.

The stretch reflexes and shortening reactions evoked in six normal subjects during reinforcement were compared with those recorded from patients with Parkinson's disease. Although the responses to stretch and shortening of triceps were similar in both groups, they differed in the biceps muscle. In Parkinson's disease increasing muscle stretch of biceps was associated with increasing reflex activity, whereas the reverse relationship occurred in the activated normal subjects. The biceps EMG during sinusoidal stretching comprised two peaks in the activated normal group, a phenomenon not seen in Parkinson's disease. It is postulated that this response in the activated normal group is due to activation or reinforcement engaging long-loop reflexes. It is concluded that the rigidity of Parkinson's disease is not simply an exaggeration of the stretch reflexes found in normal man but differs in that the effects of flexor reflex afferent nerve fibres are submerged by group Ia afferent activity, and that some long-loop reflex pathways are no longer operative in Parkinson's disease.

Electromyography↗

Modulation of the stretch reflex during volitional sinusoidal tracking in Parkinson's disease.

Sinusoidal visually-guided wrist tracking, in normal and parkinsonian subjects, was perturbed by torque transients every 90 degrees throughout the movement. Long-latency stretch reflex and volitional EMG amplitude modulations were assessed as functions of the tracking phase. Reflex modulation during tracking, both in wrist flexor and extensor muscles, was found to differ significantly between parkinsonian and normal subjects. In the parkinsonian group, the abnormality consisted of an increased reflex activity during tracking phases in which the muscle was lengthening. At these phases the reflex generated torque is opposite in direction to the volitionally generated torque and the tracking movement. No differences in the unperturbed volitional EMG modulation were observed between groups for this error constrained tracking paradigm. Significant correlations were found between ratings of bradykinesia and the amount of abnormal reflex modulation in the wrist flexor. These data suggest that a component of bradykinesia results from a defective coordination of supraspinal reflex and volitional control systems.

Aged↗

Stretch reflexes in human masseter.

The reflex response to stretch in most contracting human muscles includes both a short-latency, probably monosynaptic, excitatory component, and a longer-latency, polysynaptic excitation. However, it has been claimed that stretch of the jaw-closing muscles evokes only the short-latency response in masseter. This question was re-examined, using controlled stretches of varied rates and durations. Very brief, rapid stretches analogous to the stimuli used to investigate the 'jaw-jerk' reflex in earlier studies evoked a prominent excitatory peak in the electromyogram at monosynaptic latency excitation, but little or no longer-latency excitation. This response could be produced even by stimuli that were barely detectable by the subject. However, this prominent electrical response did not produce a measurable increase in biting force. In contrast, slower stretches evoked both a short- and a longer-latency excitatory response in the surface electromyogram, as in most limb muscles. It is shown that the absence of a long-latency excitatory response in earlier studies can be explained by the powerful reflex disfacilitation of the motoneurones that occurred at the end of the brief stretches used. Depending on the duration of the stretch, this disfacilitation is often sufficient to mask or abolish the long-latency reflex. The reflex response to stretches was not markedly affected by blocking the activation of mechanoreceptors around the teeth with local anaesthetic, indicating that receptors around the teeth cannot be playing more than a minor role in the response. The stretch-induced increase in force became greater as the velocity of the stretch decreased.

Adolescent↗

Stretch reflex responses in the human elbow joint during a voluntary movement.

1. The responsiveness of the stretch reflex is modulated during human voluntary limb movements. The influence of this modulation on the limb mechanical properties (stiffness) was investigated. 2. Subjects were taught to replicate accurately a rapid (4.0 rad s-1) targeted elbow flexion movement of 1 rad. From the onset of 12% of the trials a sinusoidal position disturbance (0.05 rad) was superimposed on the normal (trained) movement trajectory. The net joint torque (muscle torque) resisting these stretches was computed from measurements of applied torque, acceleration and limb inertia. Electromyographic (EMG) responses in the triceps brachii (TB), brachialis (Br) and biceps brachii (BB) were monitored. 3. The EMG responses to sinusoidal stretches applied early in the movement were less than those responses to perturbations applied when the arm neared the target (especially in the antagonist muscle TB). These EMG responses caused fluctuations in the resistance to the perturbation (stiffness), as described below. 4. When the perturbation frequency was low (< 4 Hz) the resistance of the elbow muscles to the stretch increased as the arm approached the target (48% increase). In contrast, when the stretch frequency was 7 Hz the resistance decreased by 63%. This decrease can be explained by the increased reflex response, since at 7 Hz the reflex response is probably timed so that it assists, rather than resists, the stretching as a result of loop delays. This reflex timing was confirmed by observing that, after abruptly stopping the sinusoidal stretch, the reflex response persisted for 100 ms and was indeed in a direction that would have reduced the resistance, had the perturbation continued. 5. The time course of the net muscle stiffness was estimated for frequencies ranging from 4 to 8 Hz and for each 40 ms interval a Nyquist plot was constructed, forming a C-shaped curve as frequency was varied. The size of this curve gave a measure of the stiffness resulting from reflex activity. When the arm neared the target this reflexive stiffness reached a maximum, and was probably comparable in size to the intrinsic (non-reflexive) muscle stiffness. Also, in four of the five subjects the viscous component of stiffness at 7 Hz dropped significantly below zero when the arm neared the target, again indicating that at this frequency the reflex was large and acted inappropriately.(ABSTRACT TRUNCATED AT 400 WORDS)

Adult↗

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.

Afferent Pathways↗

Transient reversal of the stretch reflex in human arm muscles.

1. Load perturbation responses can violate the law of reciprocal innervation between antagonist muscles under particular conditions. Thus flexor and extensor muscles of wrist and elbow joints are reflexly coactivated by the impact of a ball on the hand during a catching task. The aim of this study was to determine whether reflex coactivation can be preset within the central nervous system (CNS) or whether it is entirely due to the peripheral stimulus. To this end, we studied the behavior of stretch reflex responses of arm muscles evoked by torque motor perturbations applied before and during the catching task. 2. Subjects were instructed to catch a ball dropped from 1.6 m. A torque motor delivered perturbations to the elbow joint, resulting in angular motion at both elbow and wrist joints because of their dynamic mechanical coupling. Two series of experiments were performed that differed in the perturbation waveform. In the first series, a single torque pulse could be randomly applied at different times during the task. The corresponding responses were recovered by subtracting the average of the unperturbed trials from the averages of perturbed trials. In the second series of experiments, a train of pseudorandom pulses was applied continuously during each trial. The time-varying impulse responses were computed at 20-ms intervals by cross-correlation methods. 3. The pattern of the short-latency electromyographic responses evoked by either single pulses or pseudorandom perturbations obeyed the law of reciprocal innervation of antagonist muscles under basal conditions. However, the pattern of the responses evoked by the same perturbations around the time of ball impact on the hand consisted of a substantial coactivation of both stretched and shortening muscles. Reflex coactivation resulted from response patterns that differed at different joints. At the elbow, reflex coactivation resulted from a transient reversal of the direction of the short-latency responses of flexor muscles, with little changes of the responses of extensor muscles. At the wrist, instead, reflex coactivation resulted from simultaneous changes in the response waveform of both flexor and extensor muscles. 4. The peripheral conditions associated with the applied perturbations were constant before the time of ball impact. Thus, because the changes of the stretch reflex responses began before that time, they must have been generated within the CNS. It is here hypothesized that the reversal of the reflex responses is centrally gated by switching from the pathways of reciprocal inhibition to those of coactivation of antagonist alpha-motoneurons.(ABSTRACT TRUNCATED AT 400 WORDS)

Arm↗

Movement deficits caused by hyperexcitable stretch reflexes in spastic humans.

Spastic patients were instructed to make accurate, rapid ankle dorsiflexion and plantarflexion movements over different distances to a target. Ankle position and surface electromyograms (EMGs) from tibialis anterior (TA) and soleus (SOL) muscles were recorded. In 3 of 8 spastic patients tested, dorsiflexion evoked velocity-dependent activation of the antagonist (SOL) muscle which impeded the movement to the extent that the limb unintentionally reversed movement direction. We propose that this activation is reflex in origin since it is tightly synchronized, has a large peak amplitude, occurs about 50 ms after the initiation of the movement, and is velocity dependent. One of the 3 patients who had reflex-induced antagonist activation in dorsiflexion also demonstrated sustained clonus during plantarflexion. This usually occurred only if the target had been overshot so that the return of the limb stretched the soleus muscle and triggered clonus. We conclude that in some patients, hyperactive stretch reflexes cause movement deficits.

Adult↗

Effects of effort and EMG levels on short-latency stretch reflex modulation after varying background muscle contractions.

It is known that the short-latency stretch reflex (SLSR) is modulated by the background muscle activity when it is elicited at matched torque levels. This study was designed to examine the effects of muscle contraction types before a stretch perturbation on SLSR in the human soleus muscle (SOL) when SLSR was elicited at the same levels of effort and at matched electromyographic (EMG) activity levels. A mechanical stretch perturbation was applied to the calf muscles when the ankle joint reached a ninety degree tibio-tarsal joint angle after the muscles performed an isometric (pre-ISO), shortening (pre-SHO) and lengthening contraction (pre-LEN). Subjects were seated on an ankle ergometer chair and developed 0%, 10%, 20%, 30%, 40%, 50%, 60% and 70% ankle joint torque (AJT) of maximum voluntary isometric plantar flexion contraction at 80 degrees in pre-SHO, at 90 degrees in pre-ISO and at 100 degrees in pre-LEN. After that, isometric or dynamic contractions started, and the subjects were asked to maintain effort levels as, needed, to maintain the target torque levels until the end of the stretch. They relaxed their muscles fully after the stretch. This chain of processes was consecutively repeated 10 times. EMG signals obtained from SOL were averaged after they were high-pass filtered and full-wave rectified. Some major findings resulted: (1) there were no differences in SLSR area in the active muscle between pre-ISO and pre-SHO, whereas its waveform was steeper in pre-ISO than in pre-SHO. (2) SLSR p-to-p amplitude and waveform were larger and steeper in the active muscle than in the relaxed one in all conditions, whereas they were independent of the effort levels once the muscle was activated. This led to steady SLSR modulation in response to the background muscle contraction in the active muscle regardless of whether the SLSR was elicited at matched AJT or EMG activity levels. These findings suggest that SLSR is closely related to the muscle spindle sensitivity influenced by the following factors: (1) the background muscle contraction type, and (2) gamma motoneuron activity set by CNS based on the effort level.

Adaptation, Physiological↗

Intramuscular acupuncture-like electrical stimulation inhibits stretch reflexes in contralateral finger extensor muscles.

Electro-acupuncture is one of many physical measures used to relieve musculoskeletal pain and to improve the associated restricted range of motion. Experiments were designed to determine whether or not acupuncture-like stimulation inhibits stretch reflexes in an arm extensor muscle in human volunteers. Surface electromyographic recordings were made on the right extensor digitorum communis muscle and averaging techniques were used to study the reflex responses to brief deflection of the finger with a solenoid-driven probe. The ratio M1:M2 of two components of the reflex was reduced during continuous acupuncture-like stimulation of the contralateral first dorsal interosseus and extensor digitorum communis muscles near their motor points (acupuncture points LI 4 and LI 11). Concomitant changes in skin temperature were observed on the forehead and in the arm in which acupuncture-like stimulation was used. In control experiments, when the acupuncture needles were inserted subcutaneously and stimulated with the same current parameters at distinctly uncomfortable intensities, no change in the reflexes occurred. These findings show that acupuncture-like stimulation exerts physiologic effects on the central nervous system, mediated presumably by muscle afferent fibers. The effects may be relevant to relief of muscle spasm and musculoskeletal pain, and restoration of mobility.

Arm↗

Operant conditioning of spinal stretch reflexes in patients with spinal cord injuries.

Hyperactive spinal stretch reflexes (SSRs) often occur with spinal cord injuries (SCI). These altered SSRs may impair movement. Recent studies in monkeys and human subjects have indicated that the magnitude of SSRs can be modulated using operant conditioning. The purpose of this study was to determine whether hyperactive biceps brachii SSRs could be operantly conditioned downward. Seventeen chronic (> 1 year postlesion) spinal cord-injured patients participated. Subjects were trained to keep biceps background (prestretch) electromyographic (EMG) activity and elbow angle at predetermined levels prior to having the elbow rapidly extended by a torque motor to elicit the biceps SSR. All subjects participated in six baseline sessions over a 2-week period. Then, subjects were randomly assigned to either control or training groups for the next 24 sessions over an 8-week period. By the end of the study, training subjects had significantly reduced biceps SSRs (t test, P < 0.001), while control subjects SSRs were not significantly reduced (t test, P > 0.05). The reduced SSRs persisted for up to 4 months following cessation of training. The results of this study support the hypothesis that hyperactive SSRs can be operantly conditioned downward in SCI patients.

Adolescent↗

Stretch reflexes in human abdominal muscles.

Homonymous and heteronymous reflex connections of the abdominal muscles were investigated by the application of a tap to the muscle belly and observation of surface electromyographic responses. Reflex responses of the following abdominal muscles were investigated both ipsilateral and contralateral to the tap: rectus abdominis (RA), external oblique (EO) and internal oblique (IO). Reflexes were evoked in each of the homonymous muscles with latencies and estimated conduction velocities compatible with being evoked by Ia muscle afferents and having a monosynaptic component. Short latency heteronymous excitatory reflex connections were also observed in muscles on both ipsilateral and contralateral sides in response to the same stimulus. The latencies of the crossed responses were only marginally longer than responses evoked in the respective ipsilateral muscle. Moreover, the reflexes evoked in the IO muscle from ipsilateral and contralateral IO muscle afferents were of comparable amplitude, as were those reflexes evoked in ipsilateral and contralateral EO and RA muscles when tapping IO. These similarities in the reflex characteristics on the ipsilateral and contralateral sides suggest that abdominal muscle afferents activate similar pathways to muscles on both sides of the body. It follows that if the homonymous stretch reflex of abdominal muscles have a monosynaptic component, then a similar monosynaptic pathway activates synergistic motoneurones, not only ipsilaterally but also contralaterally.

Afferent Pathways↗

The simple frequency response of human stretch reflexes in which either short- or long-latency components predominate.

1. The stretch reflexes of the human abductor digiti minimi (ADM) and biceps brachii muscles were compared using small-amplitude sinusoidal stretching at 10-50 Hz and recording the surface EMG. The stimulus was applied either to the relevant proximal phalanx or to the biceps tendon while the muscle studied was contracting; the same amplitude was used for all frequencies (range 0.5-2 mm for ADM, 0.1-1 mm for biceps). 2. As the frequency increased, the response of ADM decreased while that of biceps increased. Neither muscle showed a minimum at 20-25 Hz, as previously found for wrist muscles and attributed to an interaction between short- and long-latency components of the reflex. 3. For both muscles, the phase of the response lagged behind the stimulus by an amount which increased approximately linearly with frequency, without the gross inflexion found for wrist muscles. Such linearity would be found for a system dominated by a fixed time delay; its value sets the slope. The slope for biceps was half that for ADM. The values of reflex delay calculated from the slope of the phase plots agreed reasonably with the absolute latencies of the responses evoked by tap or ramp stimulation. Part of the difference between the muscles was due to differences in peripheral conduction time, since ADM lies more distally. Most of it, however, was due to different reflexes being involved, with biceps being predominantly controlled by short-latency pathways and ADM by long-latency pathways. 4. For both muscles, the phase lag at any given frequency was less than that expected from the reflex latency, determined from the slope of the phase plot. Thus, sensory transduction and central transmission had produced a phase advance in the reflex. The 'neural phase advance' of biceps was appreciably larger than that of ADM, and more than would be expected from the behaviour of its spindle afferents. The excess is suggested to be due to the action of Renshaw inhibition, which ADM may lack. 5. The results were substantiated by recording from single motor units in biceps. Stretching at the present amplitudes had rather little effect on the overall rhythmic behaviour, as shown by interspike interval histograms. However, cycle histograms showed that the discharge was modulated reasonably sinusoidally by the stretching, whatever its frequency (i.e. the probability of the occurrence of a spike varied over the cycle). Cyclic changes were also found in autocorrelograms and amplitude spectra of the spike trains.(ABSTRACT TRUNCATED AT 400 WORDS)

Adult↗

Evidence for a supraspinal contribution to the human quadriceps long-latency stretch reflex.

In sitting humans a rapid unexpected lengthening of the knee extensors elicits a stretch reflex (SR) response as recorded by the electromyogram (EMG) which comprises multiple bursts. These are termed short latency responses (SLR), medium latency responses (MLR) and long latency responses (LLR). The aim of this study was to determine if a transcortical pathway contributes to any of these bursts. Flexion perturbations (amplitude =4 degrees, velocity=150 degrees/s) were imposed on the right knee joint of sitting subjects (n=11). The effect of the perturbation on the electromyographic (EMG) response of the pre-contracted quadriceps muscle to magnetic stimulation of the contralateral motor cortex was quantified. Transcranial magnetic stimulation (TMS) was applied to elicit a compound motor evoked potential (MEP) in the target muscle rectus femoris (RF), in the vastus lateralis (VL), vastus medialis (VM) and biceps femoris (BF). The MEP and SR were elicited either in combination or separately. When applied in combination the delay between the SR and the MEP varied from 0 to 150 ms in steps of 4, 5 and 10 ms. Somatosensory evoked potentials (SEPs) were recorded from four subjects during the imposed stretch to quantify the latency of the resulting afferent volley. Onset latencies of responses in RF were 25+/-2 ms for the SR and 20+/-4 ms for the MEP. The average SEP latency was 24+/-2 ms. A transcortical pathway thus has the potential to contribute to the RF SR no earlier than 54+/-6 ms (SEP + MEP + 10 ms central processing delay) following the stretch onset. The duration of the total reflex burst was 85+/-6 ms. Significant facilitation of the MEP commenced at 78 ms, coinciding with the LLR component of the stretch response. No such facilitation was observed in the synergists VL and VM, or in the antagonist BF. Our results indicate that the LLR of the RF likely involves supraspinal pathways. More importantly, of the investigated muscles, this involvement of higher centers in the shaping of the LLR is specific to the RF muscle during the investigated task.

Efferent Pathways↗

Muscle history, fusimotor activity and the human stretch reflex.

1. The previous history of contraction and length changes of a muscle influences the size of the stretch reflex and H reflex. Here we ask, is this dependence due to changes in mechanical properties of extrafusal fibres, intrafusal fibres of spindles, or both? 2. The soleus muscle of human subjects was conditioned using either a voluntary contraction or a contraction evoked by low-strength electrical stimulation, in the range 0-25 % of maximum. Following conditioning, reflexes were increased by more than twofold above the no-contraction value by a voluntary contraction of 5 % of maximum, or more, but not by electrical stimulation which presumably did not contract the intrafusal fibres of spindles. 3. When the muscle was conditioned with a contraction at a length shorter than the test length, rather than at the test length, a depressing effect on reflexes was attributed to both the burst of impulses generated in spindles when the muscle was stretched back to the test length and to a reduced stretch sensitivity of muscle spindles. 4. The experiments demonstrate the importance of keeping the muscle and its spindles in a defined mechanical state when measuring reflexes. They also point to the powerful facilitating influences of conditioning muscle contractions provided they recruit the intrafusal fibres of spindles.

Adult↗

Adaptive plasticity in the spinal stretch reflex: an accessible substrate of memory?

The study of the substrates of memory in higher vertebrates is one of the major problems of neurobiology. A simple and technically accessible experimental model is needed. Recent studies have demonstrated long-term adaptive plasticity, a form of memory, in the spinal stretch reflex (SSR). The SSR is due largely to a two-neuron monosynaptic arc, the simplest, best-defined, and most accessible pathway in the primate central nervous system (CNS). Monkeys can slowly change SSR amplitude without a change in initial muscle length or alpha motoneuron tone, when reward is made contingent on amplitude. Change occurs over weeks and months and persists for long periods. It is relatively specific to the agonist muscle and affects movement. The salient features of SSR adaptive plasticity, combined with clinical and laboratory evidence indicating spinal cord capacity for intrinsic change, suggest that SSR change eventually involves persistent segmental alteration. If this is the case, SSR plasticity should be a powerful model for studying the neuronal and synaptic substrates of memory in a primate.

Adaptation, Physiological↗