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Adapting reflexes controlling the human posture.

Doubt about the role of stretch reflexes in movement and posture control has remained in part because the questions of reflex "usefulness" and the postural "set" have not been adequately considered in the design of experimental paradigms. The intent of this study was to discover the stabilizing role of stretch reflexes acting upon the ankle musculature while human subjects performed stance tasks requiring several different postural "sets". Task specific differences of reflex function were investigated by experiments in which the role of stretch reflexes to stabilize sway doing stance could be altered to be useful, of no use, or inappropriate. Because the system has available a number of alternate inputs to posture (e.g., vestibular and visual), stretch reflex responses were in themselves not necessary to prevent a loss of balance. Nevertheless, 5 out of 12 subjects in this study used long-latency (120 msec) stretch reflexes to help reduce postural sway. Following an unexpected change in the usefulness of stretch reflexes, the 5 subjects progressively altered reflex gain during the succeeding 3-5 trials. Adaptive changes in gain were always in the sense to reduce sway, and therefore could be attenuating or facilitating the reflex response. Comparing subjects using the reflex with those not during so, stretch reflex control resulted in less swaying when the task conditions were unchanging. However, the 5 subjects using reflex controls oftentimes swayed more during the first 3-5 trials after a change, when inappropriate responses were elicited. Four patients with clinically diagnosed cerebellar deficits were studied briefly. Among the stance tasks, their performance was similar to normal in some and significantly poorer in others. Their most significant deficit appeared to be the inability to adapt long-latency reflex gain following changes in the stance task. The study concludes with a discussion of the role of stretch reflexes within a hierarchy of controls ranging from muscle stiffness up to centrally initiated responses.

Adaptation, Physiological↗

Anticipatory modulation of neck muscle reflex responses induced by mechanical perturbations of the human forehead.

The aim of this study was to test whether anticipation of upcoming head blow stimuli, which elicit reflex responses in the neck muscle, makes the reflex responses greater or not. In nine healthy subjects the reflex responses were elicited in the sternocleidomastoid (SCM) muscle in the eyes-open (EO) and eyes-closed (EC) conditions, which corresponded to the predictable and unpredictable conditions, respectively. The subjects were instructed not to resist the perturbations after the impact. The results demonstrated that the reflex response of the SCM muscle was significantly smaller in the predictable EO condition than in the unpredictable EC condition (P < 0.05), although no significant differences were observed in either the background EMG activities or the head accelerations. Further, this effect of anticipation was observed only in the later reflex EMG component, which most probably mediated the stretch reflex pathway. In contrast, no significant difference was observed in the early component, which was presumed to be the vestibular-collic reflex. The reduced stretch reflex response was suggested to be functionally relevant to the task requirement, i.e., to let the neck extension movement occur, and not to resist after the impact of the head blow. It was concluded that the anticipation has an effect on reducing the stretch reflex responses in the neck muscle, but does not have any effect on the presumed vestibular-collic reflex under the present experimental paradigm. It is suggested that the gain of the stretch reflex pathway is modulated by anticipatory information of upcoming mechanical event.

Adult↗

[Dynamic properties of the reflex to muscle stretching].

The stretch and unloading reflexes were studied on extensor muscles of the hindlimb of decerebrated cats. It had been shown that nonlinear properties of the muscle dynamics were transformed under the reflex control and hysteresis inherent in the muscle itself was not removed by the stretch reflex. Hysteresis of the stretch reflex led to uncertainty of the equilibrium value of the muscle length and the equilibrium length depended on the movement prehistory. Stiffness of the muscle lengthening was shown to depend on the point of change in the movement direction. The observed nonlinearity of the stretch reflex was discussed from the functional point of view.

Afferent Pathways↗

Segmental reflexes and ankle joint stiffness during co-contraction of antagonistic ankle muscles in man.

The size of soleus H-reflexes and short-latency stretch reflexes was measured at different levels of plantar flexion or co-contraction (simultaneous activation of dorsi- and plantar flexors) in seven healthy subjects. In four of seven subjects the short-latency stretch reflex was smaller during weak co-contraction than during isolated plantar flexion at matched background electromyogram (EMG) levels in the soleus muscle. In three of these four subjects the stretch reflex was larger during strong co-contraction than during plantar flexion, whereas it had the same size during the two tasks in the last subject. In the remaining subjects the stretch reflex either had the same size or was larger at all levels of co-contraction than at similar levels of plantar flexion. In contrast, the H-reflex was found to decrease with co-contraction at all contraction levels in all subjects. The decrease in the reflexes during weak co-contraction might be caused by presynaptic inhibition of Ia afferents. It is unclear why only the H-reflex decreased during strong co-contraction. The stiffness of the ankle joint was measured from the torque increment following the stretch of the plantar flexors divided by the stretch amplitude. In all subjects the total stiffness of the ankle joint was larger during strong co-contraction than during plantar flexion of similar strength. The stiffness was smaller during weak co-contraction than during weak plantar flexion in three out of seven subjects.(ABSTRACT TRUNCATED AT 250 WORDS)

Adult↗

Some pathophysiological aspects of spasticity and the search for rational and successful therapy.

In spite of the different uses of the term "spasticity", hyperactivity of skeletal muscle stretch reflexes is the one common factor and we therefore need to know how this is produced by lesions within the central nervous system and what are its consequences to the initiation and execution of voluntary movement, not only initially but also chronically. The alpha motoneurone is directly responsible for the initiation of skeletal muscle contraction and final integration of excitatory and inhibitory nervous input normally takes place on its surface. In spasticity there is not only loss of descending direct excitatory and inhibitory control of motoneurones, but also loss of the control of spinal interneurones which would normally regulate (principally by inhibition) segmental spinal reflexes, including the stretch reflexes, especially those concerned with antigravity muscles. Gamma motoneurones may also have a reduction inhibitory control with consequent increase of muscle spindle sensitivity to stretch, and this may be further exaggerated by changes in the physical properties of affected muscles. The peripheral disorders of function are more accessible to study and to pharmacological and physical treatment, but with the increasing knowledge of inhibitory mechanisms and their pharmacology there is hope that some degree of influence may be possible within the central nervous system, by therapy with drugs that mimic or prolong the action of inhibitory transmitters.

Animals↗

A dynamic neuromuscular model for describing the pendulum test of spasticity.

Both dynamic and static thresholds, as well as the gain in the stretch reflex loop, affect the sensitivity of motoneurons to muscle stretch. How the variation in each parameter will influence the mechanical behavior of patients with spasticity is not well understood because of the difficulty in experimentally isolating individual parameters. A neuromuscular dynamic model, based on the pendulum test of spasticity, has been developed to study the specific contribution of individual parameter abnormalities in stretch reflex loops to the observed mechanical abnormalities. The model contains detailed nonlinear dynamics of muscle force generation and stretch reflexes. A computer simulation of the model indicates that the stretch reflex thresholds and the gain have different influences on the leg swing in the pendulum test of spasticity. Individual changes in the static stretch reflex threshold, in the dynamic threshold, or in the gain can not stimulate the whole spectrum of spasticity severity. When simultaneous changes in all three parameters of the stretch reflex loop occur, a small variation of the gain coupled with changes in both static and dynamic thresholds can produce increasing severity of spasticity as the thresholds further decrease. The model is also successful in simulating the effect of posture changes on spasticity.

Adult↗

Does spasticity contribute to walking dysfunction after stroke?

OBJECTIVES: Clinically, it is assumed that spasticity of the calf muscles interferes with walking after stroke. The aim was to examine this assumption by evaluating the contribution of spasticity in the gastrocnemius muscle to walking dysfunction in an ambulant stroke population several months after stroke. METHODS: Fourteen stroke patients who were able to walk independently and 15 neurologically normal control subjects were recruited. Both resting and action stretch reflexes of the gastrocnemius muscle were investigated under conditions that simulated walking. Resting tonic stretch reflexes were measured to assess spasticity whereas action tonic stretch reflexes were measured to assess the possible contribution of spasticity to gait dysfunction. RESULTS: Two thirds of the stroke patients exhibited resting tonic stretch reflexes which indicate spasticity, whereas none of the control subjects did. However, the stroke patients exhibited action tonic stretch reflexes that were of similar magnitude to the control subjects, suggesting that their reflex activity during walking was not different from that of control subjects. Furthermore, there was no evidence that the action stretch reflex in the stroke patients contributed a higher resistance to stretch than the control subjects. CONCLUSIONS: Whereas most of the stroke patients exhibited spasticity when measured both clinically and physiologically, they did not exhibit an increase in resistance to dorsiflexion due to exaggerated action tonic stretch reflexes. It is concluded that it is unlikely that spasticity causes problems in walking after stroke in ambulant patients. Therefore, it seems inappropriate to routinely reduce or inhibit the reflex response to improve functional movement in stroke rehabilitation. Factors other than spasticity should be considered when analysing walking after stroke, so that appropriate treatment is provided to patients.

Aged↗

[Reflex examination. Neurophysiologic and biochemical status].

The neural network excites when a stretch reflex is elicited during a physical examination. It is important to be aware that the stretch reflex depends on the contraction level of the muscle before stretch, and on the amplitude and velocity of the muscle stretch. A changed resistance from passive tissue or muscle fibre them self has to be taken into account when evaluating the size of the stretch reflex. A weak contraction compared with no contraction in the stretched muscle decrease the variation in the size of the stretch reflex. Influence from other types of receptors and synergistic and antagonistic muscles can also contribute to the stretch reflex.

Animals↗

Intrinsic properties and reflex compensation in reinnervated triceps surae muscles of the cat: effect of activation level.

The manner in which activation levels influence intrinsic muscular properties and contributions of the stretch reflex were studied in homogeneous soleus (SOL) and heterogeneous gastrocnemius (G) muscles in the decerebrate cat. Intrinsic mechanical properties were represented by the initial stiffness of the muscle, measured prior to reflex action, and by the tendency of the muscle to yield during stretch in the absence of the stretch reflex. Stiffness regulation by the stretch reflex was evaluated by measuring the extent to which reflex action reduces yielding and the extent to which stiffness depends on background force. Intrinsic mechanical properties were measured in muscles deprived of effective autogenic reflexes using the method of muscular reinnervation. Reinnervated muscles were recruited to force levels comparable to those achieved during natural locomotion. As force declined during crossed-extension reflexes in reinnervated and intact muscles, initial stiffness declined according to similar convex trajectories. The data did not support the hypothesis that, for a given force level, initial stiffness is greatest in populations of predominantly type I motor units. Incremental stiffness (Deltaf/Deltal) of both G and SOL increased in the presence of the stretch reflex. Yielding of SOL (ratio of incremental to initial stiffness) substantially decreased in the presence of the stretch reflex over the full range of forces. In reflexive G, yielding significantly decreased for low to intermediate forces, whereas at higher forces, yielding was similar irrespective of the presence or absence of the stretch reflex. The stretch reflex regulates stiffness in both homogeneous and heterogeneous muscles.

Animals↗

Kinetics, Ca2+ dependence, and biophysical properties of integrin-mediated mechanical modulation of transmitter release from frog motor nerve terminals.

Neurotransmitter release from frog motor nerve terminals is strongly modulated by change in muscle length. Over the physiological range, there is an approximately 10% increase in spontaneous and evoked release per 1% muscle stretch. Because many muscle fibers do not receive suprathreshold synaptic inputs at rest length, this stretch-induced enhancement of release constitutes a strong peripheral amplifier of the spinal stretch reflex. The stretch modulation of release is inhibited by peptides that block integrin binding of natural ligands. The modulation varies linearly with length, with a delay of no more than approximately 1-2 msec and is maintained constant at the new length. Moreover, the stretch modulation persists in a zero Ca2+ Ringer and, hence, is not dependent on Ca2+ influx through stretch activated channels. Eliminating transmembrane Ca2+ gradients and buffering intraterminal Ca2+ to approximately normal resting levels does not eliminate the modulation, suggesting that it is not the result of release of Ca2+ from internal stores. Finally, changes in temperature have no detectable effect on the kinetics of stretch-induced changes in endplate potential (EPP) amplitude or miniature EPP (mEPP) frequency. We conclude, therefore, that stretch does not act via second messenger pathways or a chemical modification of molecules involved in the release pathway. Instead, there is direct mechanical modulation of release. We postulate that tension on integrins in the presynaptic membrane is transduced mechanically into changes in the position or conformation of one or more molecules involved in neurotransmitter release, altering sensitivity to Ca2+ or the equilibrium for a critical reaction leading to vesicle fusion.

Animals↗

Spastic hypertonia: mechanisms and measurement.

Spastic hypertonia has been defined as a motor disorder characterized by a velocity-dependent increase in tonic stretch reflexes (muscle tone) with exaggerated tendon jerks, resulting from hyperexcitability of the stretch reflex, as one component of the upper motor neuron syndrome. Heightened muscle tone may be the result of changes intrinsic to the muscle or to altered reflex properties. Increased motoneuronal excitability and/or enhanced stretch-evoked synaptic excitation of motoneurons are mechanisms that might enhance stretch reflexes. Two distinct parameters may be altered in the pathologic stretch reflex--the "set point," or angular threshold of the stretch reflex, and the reflex "gain," or the amount of force required to extend the limb in proportion to the increasing joint angle. Earlier studies fail to dissociate the contributions of reflex threshold and reflex gain. Recent investigations suggest that spastic hypertonia may be the result of a decrease in stretch reflex threshold without significant increase in reflex gain, as was previously believed. Various clinical scales, biomechanical paradigms, pendulum models, and electrophysiologic studies have been used to quantify spastic hypertonia. Biomechanical methods seem to correlate most closely with the clinical state. Spastic hypertonia is but one component of the upper motor neuron syndrome, whose features also include loss of dexterity, weakness, fatigability, and various reflex release phenomena. These other features of the upper motor neuron syndrome may well be more disabling to the patient than changes in muscle tone.

Humans↗

Physiology and anatomy of possible oscillators in the central nervous system.

Pathologic tremors occur when the normal, continuous pattern of muscle activation is replaced by relatively synchronous bursting. This article discusses the possible roles of stretch reflex and central oscillators in producing tremor. The gain and conduction delay in some reflex arcs places the stretch reflex close to oscillation in some muscles during normal operation. Within the brain, cells in the thalamic relay nuclei and inferior olive contain a set of ionic conductancies in the cell membrane that can interact to produce oscillations in membrane potential. This tendency is exaggerated by hyperpolarizing or depolarizing the cell away from the normal resting potential. The activity of neighboring cells can be coupled (by electrotonic gap junctions in the olive and by recurrent axonal projections from the reticular nucleus in the thalamus), thus a large population of cells can oscillate together and exert a powerful rhythmic influence on motor output.

Cerebral Cortex↗

The flexion reflex as a diagnostic tool.

Illustrated by two different cases the importance of the flexion reflex as a diagnostic tool is discussed. It is emphasised that the flexion reflex has more qualities than withdrawal in defence only. In the patient with a long-standing paraplegia a disturbance of the balance between flexion reflex and stretch reflex is of clinical importance.

Adult↗

Relief of hemiparetic spasticity by TENS is associated with improvement in reflex and voluntary motor functions.

Our previous studies showed that a single 45 min application of transcutaneous electrical nerve stimulation (TENS) prolonged soleus H and stretch reflex latencies in hemiparetic subjects. In addition, 9 daily 30 min TENS applications enhanced vibratory inhibition of the H reflex and tended to decrease hyperactive stretch reflexes. These findings suggested that longer-term TENS may be effective in reducing hemiparetic spasticity. Our present objectives were 2-fold: to determine whether longer-term repetitive TENS stimulation would lead to a reduction in clinical spasticity in hemiparetic subjects, and whether such a reduction could be associated with a decrease in stretch reflex excitability and an improvement in voluntary motor function. We compared the effects of 15 daily 60 min TENS treatments over a 3 week period, with those of placebo stimulation applied to the common peroneal nerve of the affected leg in similar groups of spastic hemiparetic subjects. Our test battery consisted of 5 measurements which assessed (1) clinical spasticity scores, (2) maximal H reflex to M response ratios, (3) vibratory inhibition of H reflex, (4) stretch reflexes, and (5) maximal voluntary isometric plantarflexion and dorsiflexion, in standing. In contrast to placebo stimulation which produced no significant effects, repeated applications of TENS over time decreased clinical spasticity (P less than 0.05), and increased vibratory inhibition of the soleus H reflex (P = 0.02) after 2 weeks. These changes occurred with a substantial improvement in voluntary dorsiflexing force up to 820%, but not plantarflexing force. They were followed by a reduction in the magnitude of stretch reflexes (P = 0.05) in the spastic ankle plantarflexor, concomitant with a decrease in the EMG co-contraction ratios after a further week of stimulation. Our results thus indicated that repeated applications of TENS can reduce clinical spasticity and improve control of reflex and motor functions in hemiparetic subjects. Furthermore, the underlying mechanisms may be due partly to an enhancement in presynaptic inhibition of the spastic plantarflexor, and partly to a possible "disinhibition" of descending voluntary commands to the paretic dorsiflexor motoneurons.

Aged↗

Increased reflex activation of the peroneus longus following application of an ankle brace declines over time.

Two experiments were performed to document the time-dependent characteristics of the peroneus longus short latency stretch reflex amplitude following application of an ankle brace. In Experiment 1, stretch reflexes were induced in 15 weightbearing subjects during an unbraced condition and braced condition. In Experiment 2, stretch reflexes were induced in 15 weightbearing subjects before and after 3 h of wearing the brace. In Experiment 1. the amplitude of the stretch reflex increased in the braced condition by about 25% relative to the non-braced condition (p = 0.006). In Experiment 2 the amplitude of the stretch reflex increased about 18% immediately after application of the brace relative to the non-braced condition (p = 0.037). After 3 h, the stretch reflex amplitude was not different from that of initial non-braced condition. Given the importance of the peroneus longus muscle in ankle complex stability, further attention should be directed to whether the increased stretch reflex gain can be exploited during rehabilitation from ankle complex injuries. The findings provide a framework by which the effect of ankle braces on ankle joint proprioception, muscle activation profiles and balance may be physiologically interpreted.

Adult↗

Masseter reflex potentials in olivo-ponto-cerebellar atrophy.

We recorded masseter reflex potentials to examine the correlation between the masseter reflex and the muscle stretch reflexes of limbs in 19 patients with olivo-ponto-cerebellar atrophy (OPCA). The patients were subdivided into hyper- (n = 5), normo- (n = 7) and hypo- (n = 7) reflexia groups according to the degrees of the conventional deep tendon jerks in the upper limbs. The masseter reflex potentials, elicited by tapping the chin with a reflex hammer, were recorded from the bilateral masseters using a pair of surface electrodes. The latency of the potentials in the hyporeflexia was significantly longer than in the other groups, while the amplitude of those in the hyperreflexia group was significantly higher than in the other groups. These results indicate that in patients with OPCA the magnitude and latency of the masseter reflex correlates with the status of the muscle stretch reflexes of the limbs in contrast with Friedreich's ataxia where the masseter reflex has been reported to be normal or hyperactive despite hyporeflexia in the limbs.

Adult↗

Reduced short and long latency reflexes during voluntary tracking movement of the human wrist joint.

In six healthy human subjects we compared changes in the strength of Hoffmann (H), short latency (30-55 ms) and long latency (55-100 ms) stretch reflexes of flexor carpi radialis (FCR) muscle during movement and isometric contractions. In one set of experiments, stretches were imposed to the wrist during voluntarily tracked sinusoidal movement and during matched isometric contractions to compare short and long latency stretch reflex responses. In the second set, H-reflexes were compared during similar matched conditions. All reflexes decreased significantly (P < 0.05) during the voluntary tracking movement. The H-reflex was reduced during the wrist flexion, on average, by 33% of its value obtained during the isometric condition. Compared with their values during isometric conditions, the short latency stretch reflex and long latency stretch reflex during movement were reduced by 52 and 40%, respectively. From the pattern changes of the stretch reflexes and the H-reflex, a movement-induced presynaptic inhibition combined with pronounced muscle spindle unloading is proposed to play an important role in decreasing the strength of the stretch reflexes during the tracking task as compared with a matched isometric contraction.

Adult↗

Influence of long-term spaceflight on neuromechanical properties of muscles in humans.

Reflex and elastic properties of the triceps surae (TS) were measured on 12 male cosmonauts 28-40 days before a 3- to 6-mo spaceflight, 2 or 3 days after return (R+2/+3) and a few days later (R+5/+6). H reflexes to electrical stimulations and T reflexes to tendon taps gave the reflex excitability at rest. Under voluntary contractions, reflex excitability was assessed by the stretch reflex, elicited by sinusoidal length perturbations. Stiffness measurements concerned the musculoarticular system in passive conditions and the musculotendinous complex in active conditions. Results indicated 1) no changes (P > 0.05) in H reflexes, whatever the day of test, and 2) increase in T reflexes (P < 0.05) by 57%, despite a decrease (P < 0.05) in musculoarticular stiffness (11%) on R+2/+3. T reflexes decreased (P < 0.05) between R+2/+3 and R+5/+6 (-21%); 3) increase in stretch reflexes (P < 0.05) on R+2/+3 by 31%, whereas it decreased (P < 0.05) between R+2/+3 and R+5/+6 (-29%). Musculotendinous stiffness was increased (P < 0.05) whatever the day of test (25%). Links between changes in reflex and stiffness were also studied by considering individual data. At R+2/+3, correlated changes between T reflexes and musculoarticular stiffness suggested that, besides central adaptive phenomena, musculoarticular structures took part in the reflex adaptation. This mechanical contribution was confirmed when data collected at R+2/+3 and R+5/+6 were used because correlations between changes in stretch reflexes and musculotendinous stiffness were improved. In conclusion, the present study shows that peripheral influences take part in reflex changes in gravitational unloaded muscles, but can only be revealed when central influences are reduced.

Elasticity↗