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Facilitation of both stretch reflex and corticospinal pathways of the tibialis anterior muscle during standing in humans.

Excitability of both stretch reflex (SR) and motor evoked potential (MEP) elicited in the tibialis anterior (TA) muscle by transcranial magnetic stimulation were tested in standing humans. The results demonstrated significantly greater values for both SR and MEP in the TA while standing than while in the supine posture, although background electromyographic activity was silent in the two conditions. Taken together with previous reports that both pathways are facilitated in the TA at the early stance phase of human walking, our findings suggest that a common neural mechanism underlies both observations, one that might be functionally relevant for securing ankle joint stabilization during upright standing.

Adult↗

Specification of synaptic connections mediating the simple stretch reflex.

A variety of mechanisms underlie the specification of synaptic connections during development. In the monosynaptic stretch reflex in vertebrates, sensory neurones innervating muscle spindles are not determined until they make contact with a particular muscle. Instead, the muscle they supply appears to specify the pattern of central connections they establish with motoneurones. Developing thoracic sensory neurones made to project to novel peripheral targets in the forelimb of tadpoles project into the brachial spinal cord, something they never do in normal frogs. Moreover, these foreign sensory neurones make monosynaptic connections with the now functionally appropriate brachial motoneurones. Normal patterns of neuronal activity are not necessary for the formation of specific central connections. Neuromuscular blockade of developing chick embryos with curare during the period of synaptogenesis does not prevent the formation of correct sensory-motor connections. Competitive interactions among the afferent fibres also do not appear to be important in this process. When the number of sensory neurones projecting to the forelimb is drastically reduced during development, each afferent fibre still makes central connections of the same strength and specificity as normal. Together, these results suggest that peripheral targets induce some molecular change in developing sensory neurones such that they can recognize their appropriate synaptic partners in the spinal cord.

Animals↗

Behaviour space of a stretch reflex model and its implications for the neural control of voluntary movement.

A nonlinear model for the stretch reflex has recently been used to study the interactions between voluntary and reflex controls during fast, targeted movements. The present study explores the topography of a 'behaviour space' generated by computer simulations of this model under various combinations of values for the gain parameters and time constants in the model's feedback loops. In general, we define a behaviour space to be any set of behavioural characteristics of the simulated movement, such as movement time, peak acceleration or peak velocity. The mathematical model can therefore be viewed as an M x N dimensional map from its parameter space N to a behaviour space M. Here, a one-dimensional behaviour space is explored. This provides a method for quantitatively comparing the different control strategies that might be employed by the nervous system for integrating reflex and descending signals during fast, voluntary movements. The results indicate that an optimal strategy will employ proprioceptive feedback as a means of fine-tuning the braking and clamping activities of fast, goal-directed movements and that descending signals are primarily important for initiating the movement and for controlling reciprocal patterns of muscle activity during the end phase of the movement.

Feedback↗

Adaptive plasticity in primate spinal stretch reflex: behavior of synergist and antagonist muscles.

Monkeys can gradually change the amplitude of the biceps spinal stretch reflex (SSR) without change in initial muscle length or biceps background electromyographic activity (EMG) (17). We investigated the concurrent behavior of synergist (brachialis and brachioradialis) and antagonist (triceps) muscles. Synergist background EMG remained stable while marked change occurred in biceps SSR amplitude. Triceps background EMG was minimal under all conditions. Thus biceps SSR amplitude change was not due to change in the background activity of closely related muscles. When biceps SSR amplitude changed, synergist SSR amplitude changed similarly but to a lesser extent. Brachialis change averaged 72% of biceps change, while brachioradialis change averaged 33%. By indicating that SSR amplitude change is relatively specific to the agonist muscle, this finding eliminates a number of nonspecific mechanisms as possible origins of SSR amplitude change. Thus it supports the potential value of the SSR as a system for studying the neuronal and synaptic bases of memory in the primate central nervous system (CNS).

Adaptation, Physiological↗

Effects of dorsal cord stimulation on stretch reflexes.

The effects of dorsal cord stimulation on phasic and tonic stretch reflex activity in extensor muscles were studied in decerebrate cats. The tonic stretch response was depressed in both fore- and hindlimb (stimulation at levels C1 and T8, respectively) and this often persisted for 5-20 min after the end of 1-10 min of dorsal cord stimulation. Depression of the phasic stretch response was only consistently seen in the forelimb during stimulation (C1) and this rarely outlasted the period of stimulation. These results support the idea that dorsal cord stimulation can reduce muscle tone but provide no explanation for the long-lasting effects of chronic, continuous stimulation in spastic man.

Animals↗

Contralateral and long latency effects of human biceps brachii stretch reflex conditioning.

Results from previous studies on monkeys and human subjects have demonstrated that the biceps brachii spinal stretch reflex (SSR) can be operantly conditioned. The extent to which conditioning paradigms influence contralateral SSRs or longer latency responses in the same limb has not been examined. Nine subjects were given 10 training sessions to either increase or decrease the size of their biceps brachii SSR. Group changes were compared to the mean of six baseline (control) sessions. Both groups showed progressive SSR changes over the training sessions. Up-trained subjects increased their SSR responses by an average of 135.3% above baseline, with the last three sessions showing a 237.5% increase, while down-trained subjects reduced their average SSR responses by 43.4%, with a 52.7% reduction over the last three sessions. ipsilateral longer latency responses showed average changes of 68.9% and -68.7% for up- and down-trainers, respectively. As in the case of SSRs, these responses changed progressively over sessions, with a 131.5% increase seen in the last three up-training sessions and an 82.4% reduction over the same period for down-trainers. Correlation coefficients between SSR and longer latency responses were high (R = 0.90, up-trainers; R = 0.87, down-trainers). Contralateral SSR and longer latency responses, measured in the absence of feedback and at least 10 min after ipsilateral conditioning, showed directional changes that were similar to the trained side, but their magnitudes were not as profound. Collectively, these data suggest that unilateral SSR conditioning affects spinal circuits controlling contralateral SSRs and influences longer latency responses.

Adult↗

Improved organotypic cell culture model for analysis of the neuronal circuit involved in the monosynaptic stretch reflex.

Knowledge regarding neuronal circuit formation is central for the understanding of the vast network making up the brain. It is therefore necessary to find novel ways to analyze the mechanisms involved in well-defined neural circuits. We present an improved in vitro model of the monosynaptic stretch reflex circuit, based on primary organotypic cell cultures. By using limb tissue as a source of muscle fibers instead of circumspinal tissue we could make the in vitro system more in vivo like in the sense that it focuses on the stretch reflex involving limb muscles. Furthermore, our analyses showed that this procedure allows muscle fibers to follow the normal developmental pattern. Particularly interesting was the finding of slow tonic myosin heavy chain expressing muscle fibers, a developmental marker for muscle spindles, in the cultures showing that this system has the potential to contain the complete reflex circuits.

Animals↗

After-effects on stiffness and stretch reflexes of human finger flexor muscles attributed to muscle thixotropy.

1. While the subject maintained a weak contraction in his finger flexor muscles, holding the metacarpophalangeal joints in 45 deg flexion, test torque pulses were applied which caused rapid finger extension movements and electromyographic (EMG) stretch reflex responses. Before each test pulse the fingers were passively flexed or extended ('post-short' and 'post-long' trials) for about 10 s. The EMG and joint deflection responses in the two types of trial were compared after averaging. 2. In the 'post-long' trials, the EMG reflex response showed a comparative increase in latency, with a reduction of the short-latency (M1) component and an enhancement of the medium-latency (M2) component. 3. The angular deflections were larger, and the turning points of the deflections, which indicated the start of the mechanical reflex responses, occurred later in the 'post-long' trials. These differences were not seen when the torque pulse was immediately preceded by a strong, brief isometric finger flexor contraction in the test position. 4. Immediately following the return to the test position the background finger flexor EMG activity was larger in the 'post-long' trials, a difference which gradually subsided over 15-20 s. A strong, brief contraction in the test position also eliminated this inter-trial difference. 5. The results are interpreted as manifestations of thixotropic after-effects in intra- and extrafusal muscle fibres. It is proposed that the M1 component of the stretch reflex is largely a response to the 'initial burst' of impulses in primary spindle afferents.(ABSTRACT TRUNCATED AT 250 WORDS)

Adult↗

Temporal facilitation of spastic stretch reflexes following human spinal cord injury.

Recent evidence suggests that alterations in ionic conductances in spinal motoneurones, specifically the manifestation of persistent inward currents, may be partly responsible for the appearance of hyperexcitable reflexes following spinal cord injury (SCI). We hypothesized that such alterations would manifest as temporal facilitation of stretch reflexes in human SCI. Controlled, triangular wave, ankle joint rotations applied at variable velocities (30-120 deg s(-1)) and intervals between stretches (0.25-5.0 s) were performed on 14 SCI subjects with velocity-dependent, hyperexcitable plantarflexors. Repeated stretch elicited significant increases in plantarflexion torques and electromyographic (EMG) activity from the soleus (SOL) and medial gastrocnemius (MG). At higher velocities (> or = 90 deg s(-1)), reflex torques declined initially, but subsequently increased to levels exceeding the initial response, while mean EMG responses increased throughout the joint perturbations. At lower velocities (< or = 60 deg s(-1)), both joint torques and EMGs increased gradually. Throughout a range of angular velocities, reflex responses increased significantly only at intervals < or = 1 s between stretches and following at least four rotations. Ramp-and-hold perturbations used to elicit tonic stretch reflexes revealed significantly prolonged EMG responses following one or two triangular stretches, as compared to single ramp-and-hold excursions. Post hoc analyses revealed reduced reflex facilitation in subjects using baclofen to control spastic behaviours. Evidence of stretch reflex facilitation post-SCI may reflect changes in underlying neuronal properties and provide insight into the mechanisms underlying spastic reflexes.

Adult↗

Comparison of stretch reflex activities and reaction times in two separate age groups of human subjects.

The effect of prior instruction on stretch reflex activity was compared in two separate age groups of healthy subjects. No clear differences were found for amplitude modulation and reaction items between the younger (25.6 +/- 3.2 years) and the older group (56.6 +/- 6.5 years) of subjects. A significant difference was observed in the absolute magnitudes of the background activity, short latency and long latency reflex activities which were higher in the older group of subjects than in the younger subjects.

Adult↗

Effects of arousal and natural baroreceptor activation on the human muscle stretch reflex.

The nociceptive flexion reflex is inhibited during systole; this inhibition may be due to increased baroreceptor stimulation. It is yet to be determined whether other spinal reflexes are similarly modulated across the cardiac cycle. There is also evidence that stretch and tendon reflexes are facilitated by increased arousal. This study investigated the effects of phase of the cardiac cycle and arousal on the muscle stretch reflex components M1, M2, and M3. Stretch reflexes were elicited in leg muscles at six intervals across the cardiac cycle during rest, number repetition, and mental arithmetic. Mental arithmetic provoked increased cardiovascular arousal and facilitated both M1 and M2 compared to rest and number repetition. The stretch reflex did not vary with the phase of the cardiac cycle. While the stretch reflex is susceptible to arousal, natural baroreceptor-mediated modulation across the cardiac cycle may be specific to nociception.

Adult↗

Recruitment order of motor units during the stretch reflex in man.

The recruitment pattern of single motor units contributing to the short- and long-latency periods of the stretch reflex was investigated in human adults. When non-tonic, a given unit responded to stretch during the long-latency reflex period. When made tonic, the same unit showed a reflex response during the short-latency period, while a higher threshold, non-tonic unit would respond during the long-latency period. The data show an orderly recruitment of motor units during both phases of the stretch reflex in man.

Adult↗

Time-course analysis of stretch reflexes in hemiparetic subjects using an on-line spasticity measurement system.

Spasticity after a stroke is usually assessed in a score form by subjectively determining the resistance of a joint to an externally imposed passive movement. This work presents a spasticity measurement system for on-line quantifying the stretch reflex of paretic limbs. Four different constant stretch velocities in a ramp-and-hold mode are used to elicit the stretch reflex of the elbow joint in spastic subjects. The subjects are tested at supine position with the upper limb stretched towards the ground, in contrast with the horizontally stretched movement used in other studies. By subtracting the baseline torque, reflex torque measured at a selected low stretch velocity of 5 deg/sec, the influence of gravity torque and inertial in vertical stretching mode can be minimized. The averaged speed-dependent reflex torque (ASRT), defined as the measured torque deviated from the baseline torque, is used for quantifying the spastic hypertonia. Four subjects having incurred cerebrovascular accident (CVA) are recruited for time-course study in which the measurements are taken at 72 hours, one week, one month, three months, and six months after onset of stroke. During the development of spasticity, the changes of ASRT and velocity sensitivity of ASRT of the involved and the intact elbow joints are discussed.

Elbow Joint↗

Stretch reflex sensitivity: effects of postural and muscle length changes.

In this study, a combination of clinical evaluation, laboratory testing, and model simulation of spasticity is performed under various postural conditions to investigate the changes in the sensitivity and specific mechanisms of spasticity. Fifty-nine multiple sclerosis patients participated in the study and received spasticity evaluation based on both the Ashworth scale and the pendulum test. Spasticity was found to increase in the pendulum test when the subjects were tested in a supine posture, compared to when they were sitting. Three patterns of stretch reflex response were seen for similar leg swing trajectories. While it was clear that the increased stretch of rectus femoris in the supine posture contributed to the increased spasticity, the results of modeling showed that other more complex factors were also involved. The supraspinal descending modulation associated with postural control may play a more dominant role in the severity of spasticity. The results suggest that the biomechanical test of spasticity should be performed for several different postures or ranges of movement with muscle activities monitored simultaneously, so that the effect of various factors can be examined. The work also indicates that a neuromusculoskeletal model with detailed muscle dynamics and stretch reflex loops is a valuable tool for investigating the neural mechanisms of spasticity.

Adult↗

Trigeminal mesencephalic neurons innervating functionally identified muscle spindles and involved in the monosynaptic stretch reflex of the lateral pterygoid muscle of the guinea pig.

Location of the neurons in the trigeminal mesencephalic nucleus innervating stretch receptors of the lateral pterygoid muscle and the mode of their synaptic connection on the lateral pterygoid motoneurons of the guinea pig were studied physiologically as well as morphologically, in comparison with the trigeminal mesencephalic neurons innervating muscle spindles in the superficial masseter muscle, with the following results: stimulation of the caudal half of the trigeminal mesencephalic nucleus evoked monosynaptic excitatory postsynaptic potentials in the ipsilateral lateral pterygoid motoneurons. Stimulation of the lateral pterygoid nerve directly evoked spike potentials in the neurons located in the caudal half of the ipsilateral trigeminal mesencephalic nucleus, which responded with increased firing to stretch, and with silent period to twitch, of the ipsilateral lateral pterygoid muscle. Averaging of intracellular potentials of the lateral pterygoid motoneurons with extracellular spike potentials of these trigeminal mesencephalic neurons revealed excitatory postsynaptic potentials after a monosynaptic latency, but no inhibitory postsynaptic potentials. Injection of horseradish peroxidase into the lateral pterygoid muscle labeled 15-20 cells in the caudal half of the ipsilateral trigeminal mesencephalic nucleus, while 174-228 cells retrogradely labeled by horseradish peroxidase were found throughout the whole rostrocaudal extent of the ipsilateral trigeminal mesencephalic nucleus following injection of horseradish peroxidase into the masseter muscle. It was concluded that neurons in the caudal half of the trigeminal mesencephalic nucleus send their peripheral processes to stretch receptors, presumably muscle spindles, in the ipsilateral lateral pterygoid muscle and that their central processes have excitatory synapses on ipsilateral lateral pterygoid motoneurons, thus comprising the afferent limb of a monosynaptic stretch reflex arc of the lateral pterygoid muscle of the guinea pig.

Animals↗

Stretch reflex modulation during a cyclic elbow movement.

Small torque pulses were delivered to the forearm in order to test the stretch reflex of the brachialis and triceps arm muscles in 11 normal subjects performing a cyclic movement about the elbow in the horizontal plane. The flexion-extension movement was paced by a metronome and performed under various loading conditions. Reflexes for each muscle were tested either in each 50 msec segment of the 2 sec cycle period, or in a smaller number of selected phases. A late reflex, appearing at a latency of about 60 msec (measured from the onset of the torque increment), was modulated extensively during the movement cycle. The amplitude of the late reflex increased markedly at the onset of a muscle contraction. In many of the subjects reflex responsiveness began to increase as early as 200 msec prior to the onset of voluntary muscle activity. Peak reflex responses were elicited by stimuli delivered 100-150 msec prior to the peak rate of increase of dynamic load (composed of inertial, viscous and elastic forces). The increase in responsiveness was followed by a drop which was generally coincident in time with the peak rate of increase of the load opposing muscle contraction. The modulation of the late reflex is appropriately timed for reflex-generated tension to help counteract dynamic loads, intrinsic to the movement.

Adult↗

Relation between changes in long-latency stretch reflexes and muscle stiffness in Parkinson's disease--comparison before and after unilateral pallidotomy.

OBJECTIVE: To study the effects of posteroventral pallidotomy on both the size of long-latency stretch reflex (LLR) and the muscle stiffness in the wrist flexor muscles. PATIENTS AND METHODS: Eleven consecutive patients (right-handed, 6 men and 5 women) underwent left-side microelectrode-guided pallidotomy. The LLR of the contralesional forearm was studied at baseline and 2-3 months after surgery while patients continued to take their optimal medical regimens ('on' period). Patients were instructed not to respond to the perturbation (passive mode) or to oppose the mechanical extensor perturbation (active mode). RESULTS: The stretch reflex evoked by extension perturbations of the wrist consisted of a short-latency reflex (M1) and an LLR (M2). Pallidotomy had no effects on the size of M1 components in both passive and active mode and on that of M2 component in the passive mode, however, it significantly reduced M2 component in the active mode (P<0.05). The inherent muscle stiffness over 60 ms period of mechanical stretch was not influenced by pallidotomy in any experimental condition (preoperative vs. postoperative or passive mode vs. active mode). The hand displacement following M2 component increased significantly after pallidotomy in both passive (P<0.005) and active mode (P<0.05). The inverted value of the displacement following M2 component correlated with the size of M2 component (r=0.60, P<0.001). CONCLUSIONS: Pallidotomy decreased the transcortical reflex gain, probably at cortical level, and consequently reduced muscle stiffness.

Adult↗

Disturbed modulation of the stretch reflex gain during standing in cerebellar ataxia.

The postural control disturbance in patients with spino-cerebellar degeneration (SCD) was evaluated by the displacement of the center of foot pressure (CFP) and the amplitude of the triceps surae muscle H reflex in various standing postures. Twelve patients and 8 age-matched normal subjects (NL) were studied. The CFP, surface electromyograms (EMG) in the lower leg and the soleus H reflex were recorded in all subjects. The CFP displacement area was recorded in 10 patients. The following results were obtained: (1) the range of displacement of CFP from forward to backward leaning as a fraction of foot length was significantly smaller in the patients than in the controls (mean +/- S.D.; SCD = 52.8 +/- 12.7%; NL = 62.3 +/- 9.9%; P less than 0.05%, Student's t test); (2) the ratio of the increase in amplitude of the H reflex to soleus muscle EMG activity when the subjects leaned forward from an upright posture was significantly higher in the patients than in the controls (SCD = 0.41 +/- 0.40; NL = 0.17 +/- 0.15, P less than 0.05); (3) this ratio correlated with the area of CFP displacement during upright standing in the patients (r = 0.75, n = 10, P less than 0.05). These results indicate that the suppression of the stretch reflex observed in normal controls during standing is impaired in patients with SCD. We conclude that a decreased suppression of the stretch reflex is one of the mechanisms responsible for the instability on standing in patients with SCD.

Adult↗