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Sensitivity of H-reflexes and stretch reflexes to presynaptic inhibition in humans.

The sensitivity of soleus H-reflexes, T-reflexes, and short-latency stretch reflexes (M1) to presynaptic inhibition evoked by a weak tap applied to the biceps femoris tendon or stimulation of the common peroneal nerve (CPN) was compared in 17 healthy human subjects. The H-reflex was strongly depressed for a period lasting up to 300-400 ms (depression to 48 +/- 23%, mean +/- SD, of control at a conditioning test interval of 70 ms) by the biceps femoris tendon tap. In contrast, the short-latency soleus stretch reflex elicited by a quick passive dorsiflexion of the ankle joint was not depressed. The soleus T-reflex elicited by an Achilles tendon tap was only weakly depressed (92 +/- 8%). The H-reflex was also significantly more depressed than the T-reflex at long intervals (>15 ms) after stimulation of CPN (H-reflex 63 +/- 14%, T-reflex 91 +/- 13%; P < 0. 01). However, the short-latency (2 ms) disynaptic reciprocal Ia inhibition evoked by stimulation of CPN was equally strong for H- and T-reflexes (H-reflex 72 +/- 10%, T-reflex 67 +/- 13%; P = 0.07). Peaks in the poststimulus time histogram (PSTH) of the discharge probability of single soleus motor units (n = 53) elicited by an Achilles tendon tap had a longer duration than peaks evoked by electrical stimulation of the tibial nerve (on average 5.0 ms as compared with 2.7 ms). All parts of the electrically evoked peaks were depressed by the conditioning biceps femoris tendon tap (average depression to 55 +/- 27% of control; P < 0.001). A similar depression was observed for the initial 2 ms of the peaks evoked by the Achilles tendon tap (69 +/- 48%; P < 0.001), but the last 2 ms were not depressed. Conditioning stimulation of the CPN at long intervals (>15 ms) also depressed all parts of the electrically evoked PSTH peaks (n = 34; average 65%; P < 0.001) but had only a significant effect on the initial 2 ms of the peaks evoked by the Achilles tendon tap (85%; P < 0.001). We suggest that the different sensitivity of mechanically and electrically evoked reflexes to presynaptic inhibition is caused by a difference in the shape and composition of the excitatory postsynaptic potentials underlying the two reflexes. This difference may be explained by a different composition and/or temporal dispersion of the afferent volleys evoked by electrical and mechanical stimuli. We conclude that it is not straightforward to predict the modulation of stretch reflexes based on observations of H-reflex modulation.

Adult↗

Does spasticity result from hyperactive stretch reflexes? Preliminary findings from a stretch reflex characterization study.

PURPOSE: To characterize the stretch reflex response of the biceps brachii in stroke patients with elbow spasticity (prior to or within 15 min of treatment with botulinum toxin) and non-impaired volunteers with the aim of quantifying the stretch reflex excitability and observe the differences between the groups. METHODS: A cross-sectional study. Stretch reflexes from the biceps brachii were elicited following a controlled elbow extension. The amplitude, latency, rise time and duration, calculated from surface EMG recordings from the biceps brachii, were used to characterize the stretch reflex response. RESULTS: Seventeen non-impaired and 14 stroke patients participated. The amplitude was significantly lower in stroke patients than in non-impaired volunteers (p<0.05). The latency was significantly shorter in stroke patients than in non-impaired volunteers (p<0.05). There were no significant differences in rise time or duration (p>0.10). DISCUSSION: Reduction in the amplitude in stroke patients was unexpected suggesting the stretch reflex is not necessarily hyper-excitable in people with clinically diagnosed spasticity. Latency differences suggest decreased presynaptic inhibition and/or increased motor neurone excitability can occur following a stroke. However, carry over effects from previous botulinum toxin treatment may have confounded amplitude measurements. Further work evaluating the excitability of the stretch reflex independent of Botulinum toxin and its contribution to resistance to passive stretching is being conducted.

Adult↗

Neurophysiological methods for the assessment of spasticity: the Hoffmann reflex, the tendon reflex, and the stretch reflex.

PURPOSE: To review the literature concerning neurophysiological methods to assess spasticity with respect to mechanisms and methodology, and to describe the three most commonly used methods: the Hoffmann reflex (H-reflex), the Tendon reflex (T-reflex), and the Stretch Reflex (SR). METHOD: A systematic internet database search was performed to identify neurophysiological measurement methods of spasticity. A systematic exclusion procedure resulted in 185 included references, completed by additional informal search. For this paper, information about the H-, T- and stretch reflexes was extracted from these references. RESULTS: Although the reflexes are basically monosynaptic, there are many supraspinal pathways which modulate the responses in terms of their amplitude and latency. As a consequence the methods are sensitive to a considerable number of experimental conditions and are characterized by a moderate reliability and sensitivity. Correlations with other (i.e. biomechanical, neurophysiological or clinical) spasticity assessment parameters are moderate to poor. Standardised and broadly accepted protocols are still largely lacking preventing an effective exchange of knowledge. CONCLUSIONS: The clinical and experimental use of the three methods is restricted due to moderate reliability and sensitivity. It is recommended to perform combined neurophysiological-biomechanical assessment of spasticity during active, functional movement.

Electromyography↗

Interaction between short- and long-latency components of the human stretch reflex during sinusoidal stretching.

1. Using wrist muscles, the subdivision of the human stretch reflex into separate components was examined with small amplitude sinusoidal stretching of relatively high frequency (10-40 Hz). The reflex was evoked by angular rotation of the wrist (below 1 deg amplitude), applied via the hand during maintained voluntary contraction of the muscle studied; both flexors and extensors were tested. 2. The reflex response was recorded electromyographically rather than mechanically. For each condition, the surface EMG was rectified and averaged to give a cycle average showing the mean response evoked by a cycle of stretching. The cycle average was fitted with a sinusoid, the amplitude and phase of which were used to assess the reflex; their value reflects the combined action of its various sub-components. Fourier analysis gave similar results (the EMG was then rectified but not averaged). 3. The amplitude of the reflex response typically fell to a minimum in the region of 20-25 Hz. The phase lag of the response in relation to the stimulus increased approximately linearly with frequency, except in the region of the amplitude minimum. Here the lag tended to remain constant, or to decrease slightly; this created a discontinuity between the upper and lower limbs of the phase plot. 4. Such effects are attributed to an interaction between two components of the reflex that differ in latency by 20-25 ms. These would progressively change their relative phase as the frequency increased. At first they would come to interfere with each other, but then the more delayed reflex produced by a given cycle of stretching would begin to sum with the shorter latency reflex evoked by the next stretch. 5. At high and low frequencies the cycle averages were normally well fitted by a single sinusoid. Around 20-25 Hz, however, they typically showed appreciable harmonic distortion, with the second harmonic larger than the fundamental. The cycle average then showed two separate responses per cycle of stretching. These were considered to represent the uncancelled non-linear residua of separate components of the reflex response. Their relative timing shifted appropriately with change of frequency. 6. These 'double responses' are unlikely to be due to mechanical resonance. First, the relative sizes of the two components could be altered by the reflex action of cutaneous afferents. Second, the same pattern of behaviour was found when the mechanical stimuli were applied directly to the tendon of flexor carpi radialis while the hand remained fixed.(ABSTRACT TRUNCATED AT 400 WORDS)

Electromyography↗

Pulsatile motor output in human finger movements is not dependent on the stretch reflex.

1. Stretch perturbations were delivered during slow voluntary finger movements with the aim of exploring the role of the stretch reflex in generating the 8-10 Hz discontinuities that characterize these movements. Afferent activity from muscle spindle primary endings in the finger extensor muscles was recorded from the radial nerve, along with the EMG activity of these muscles, and kinematics of the relevant metacarpo-phalangeal joint. 2. Perturbations elicited a distinct response from the muscle spindles appearing at the recording electrode after 13 ms, and weak reflex responses from the muscle with peak values at 53 and 63 ms during flexion and extension, respectively. 3. The time relations between kinematics, spindle firing and modulations of EMG activity elicited by the perturbations were compared with those of the self-generated discontinuities. These analyses indicate that stretch reflex mechanisms cannot account for the modulations of EMG activity that give rise to successive 8-10 Hz discontinuities. 4. A comparison of the reflex responses to perturbations with the EMG modulations during self-generated movements indicates that the reflex was too weak to account for the pulsatile motor output during voluntary movements. 5. By inference it was concluded that the 8-10 Hz discontinuities during self-generated movements are probably generated by mechanisms within the central nervous system.

Adult↗

On the stability of delay equation models of simple human stretch reflexes.

Human stretch reflexes (SRs) are often too weak and ineffectual to provide adequate postural regulation or rhythmic movement boosting (e.g. in ankle pushoff at the end of stance phase in fast running). Recent improvements in methods of artificially enhancing skeletomotor responses, especially in therapeutic regimens, should not be widely employed until the clonus-resisting stability properties of SRs are better understood. We formulate an idealized linear servo model of a segmentally-mediated SR system which includes the often ignored electromechanical coupling delay. For typical closed-loop (delay/gain) ratios, the model is shown to be unstable for all values of loop gain when operating as a position servo, but maximally stable when operating as a velocity servo. We claim that the velocity servo or more of its nonlinear relatives is a better model for some well studied SRs than, e.g., Houk's stiff muscle hypothesis. We also present evidence that even feeble and quickly saturating monosynaptic postural servos are always unstable if operated as pure position regulators.

Humans↗

Improvement in linearity and regulation of stiffness that results from actions of stretch reflex.

We studied stretch reflexes of soleus muscles of intercollicularly decerebrated cats using a new technique for estimating the component of a stretch reflex that results from the purely mechanical properties of the active muscle (mechanical response). The difference between a net stretch reflex and its underlying mechanical response provided a direct measure of reflex action. 1. The relative contributions of reflex action and the mechanical response are different for stretch and release. With stretch, reflex action is generally large and the mechanical response small. The opposite is true with release. The property that remains relatively constant, when stretch and release are compared, is the net stiffness in opposition to length change. We concluded that reflex action compensates for variations in the inherent stiffness of the muscle. 2. Compensation is effective over a range of intermediate values of initial force, but it fails whenever the reflex force approaches zero or the maximal force at that length. 3. Reflex action is capable of modifying muscular force within 22 ms of the onset of length change. This indicates that even during a fast gallop, there is sufficient time for reflex action. 4. The mechanical properties of the active soleus muscle are highly nonlinear; e.g., muscular stiffness becomes negative transiently during stretch. In stable decerebrate preparations we found that reflex action resulted in a considerable improvement in linearity. 5. Our results support two complementary hypotheses: a) stiffness may be the regulated property of the stretch reflex, and b) the main function of autogenetic reflexes may be to conpensate for variations in the properties of skeletal muscle rather than to oppose changes in load.

Animals↗

An investigation of the servobehavior of the stretch reflex.

The stretch reflex is often described as a device for assisting in the regulation of muscle length or as a spinal servomechanism. Observation of the response to experimental interruption of voluntary movements fails to demonstrate a significant role for spinal reflexes. Functional responses with a latency of 120 ms, implying supraspinal mediation, are observed.

Electromyography↗

Reduced stretch-reflex sensitivity after exhausting stretch-shortening cycle exercise.

The stretch-shortening cycle (SSC) is an effective and natural form of muscle function but, when repeated with sufficient intensity or duration, it may lead to muscle damage and functional defects. A reduced tolerance to impact has been reported, which may be partly attributed to a reduced stretch-reflex potentiation. The aim of the present study was to examine the influence of SSC-induced metabolic fatigue and muscle damage on the efficacy of stretch reflexes, as judged by the electromyograph (EMG) response of two shank muscles (lateral gastrocnemius LG, soleus SOL) to controlled ramp stretches. These EMG responses were recorded before and immediately after exhausting SSC-type leg exercise and 2 h, 2 days and 4 days later. Serum concentrations of creatine kinase ([CK]), myoglobin and lactate were measured repetitively along the protocol. Two maximal vertical drop jumps and counter-movement jumps were performed after each reflex test. The exhausting SSC-type exercise induced an immediate reduction (P < 0.05) with a delayed short-term recovery of the LG peak-to-peak reflex amplitude. This was not accompanied by significant changes in the reflex latency. The drop jump performance remained slightly but significantly reduced (P < 0.05) until the 2nd day postexercise. Peak [CK] appeared for all the subjects on the 2nd day, suggesting the presence of muscle damage. The increase in [CK] between the 2nd h and the 2nd day postexercise was found to be negatively related (P < 0.001) to the relative changes in the drop jump height. Furthermore, a significant relationship (P < 0.05) was found between recovery of the stretch reflex in LG and the decrease of [CK] between the 2nd and the 4th day. These findings support the hypothesis of a reduced stretch-reflex sensitivity. While the exact mechanisms of the reflex inhibition remain unclear, it is emphasized that the delayed recovery of the reflex sensitivity could have resulted from the progressive inflammation that develops in cases of muscle damage.

Adult↗

Regulatory actions of human stretch reflex.

1. The stretch reflex in the elbow flexor musculature was studied in 23 human subjects. The subjects were required to establish an initial force equivalent to 10% maximum at a prescribed initial length; mechanical disturbances delivered at random times increased load force to 15% or reduced it to 5%. We measured arm force, displacement, and EMG (usually biceps); acceleration was calculated from displacement, and average responses from sets of 10 like trials. 2. Modification of the stretch reflex was studied by comparing average responses obtained with different instructions, but with the same disturbance. The usual introductions were "compensate for arm deflection" and "do not intervene voluntarily". The initial response did not depend on instruction; changes in response that depended on instruction began abruptly after a latent period which ranged from 70 to 320 ms (measured from force and acceleration), depending on conditions and subject. The latency became longer (10-50 ms) and more variable when the subject did not know the direction of disturbance in advance. This and other observations indicate that modifications of the stretch reflex are not produced by servo actions. They are produced by triggered reactions, which occur at both short and long latencies and which have properties resembling the movements produced in a reaction-time task. 3. We confirmed that most subjects can suppress triggered reactions when the instruction calls for no intervention, leaving an unmodified reflex response. This response consists of a compliant deflection of the arm in the direction of the disturbance. 4. The compensatory actions associated with unmodified stretch (and unloading) reflexes were assessed from EMG responses of biceps. During a 300-ms transient phase, EMG changes were notably asymmetric when responses to symmetric disturbances were compared. Increased force stretched biceps and produced a prominent increase in EMG, whereas decreased force allowed biceps to shorten and produced either an EMG decrease of smaller magnitude or an actual increase. These asymmetric reflex actions produced quite symmetric mechanical responses (arm displacements and forces), which implies the existence of and compensation for nonlinear muscle mechanical properties. This result is discussed in relation to the hypothesis that the function of the stretch reflex is to compensate for variations in muscle properties, thus maintaining stiffness. 5. Effective control of muscle length or joint position does not result from servo action by the stretch reflex. Errors in position are corrected only when triggered reactions are superimposed on the reflex response.

Adult↗

Development of the stretch reflex in the newborn: reciprocal excitation and reflex irradiation.

The stretch reflex is a spinal cord pathway between a muscle's stretch receptors and its own motor units. This reflex is thought to play an important role in normal motor function, because it is unique as a monosynaptic pathway, and because its hyperactivity is a hallmark of many motor disorders. We previously reported a difference in the stretch reflexes in healthy newborn infants and nonambulatory individuals with cerebral palsy (CP): these reflexes are characterized by responses from the stretched muscle and from the reciprocal or antagonist muscle. We proposed the existence of a functional spinal cord pathway that connects both agonist and antagonist muscles at a single joint. We hypothesized that this "reciprocal excitation" is a functional pathway of all newborn infants, which during the normal course of development of motor skills in infants is eliminated. If the CNS is damaged at birth, as in CP, the pathway of reciprocal excitation endures and is reinforced. In the current study of newborns, we recorded stretch reflex responses from all major muscle groups of the lower limb. This "irradiation of reflexes" is a normal phenomenon of the newborn CNS. This pathway becomes suppressed during normal maturation and control of coordinated limb movements.

Achilles Tendon↗

Development of a quantitative reflex hammer for measurement of tendon stretch reflex.

Quantification of tendon stretch reflex requires precise measurement of the tapping force of a reflex hammer. A quantitative reflex (QR) hammer consisting of two cut rubber pieces from a generic rubber reflex hammer and a uniaxial force transducer was constructed. Finite element stress analyses were conducted to estimate the natural frequency characteristics of the hammer and to find the stress distributions during the impact. Pendulum impact testing was conducted at four different heights to assess the calibration linearity and repeatability of the measurement. The QR hammer had a fundamental natural frequency of 515 Hz and showed minimal displacement and stress at the tip from the finite element simulation of the impact. The QR hammer also provided reliable and repeatable measurements as demonstrated with high coefficients of determination, exceeding 0.994 and small coefficients of variations, less than 4%. The calibration linearity was 0.64% compared with the reference force platform measurement. The QR hammer demonstrated sufficient accuracy and reliability for precise clinical assessment of tendon stretch reflexes.

Calibration↗

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.

Animals↗

Dependence of stretch reflexes on amplitude and bandwidth of stretch in human wrist muscle.

The tonic stretch reflex was investigated using small-amplitude displacements (<4.2 degrees ) of the wrist while subjects maintained average contraction levels of 25% of maximum in flexor carpi radialis. The wrist displacements were designed to preclude voluntary following but at the same time were confined to the frequency range most relevant to voluntary movements. They included a broad-frequency band (0-12 Hz) signal as well as sets of narrow-band signals spanning the range from 0 to 10 Hz. The maximum frequency was set so as to remain within the linear encoding bandwidth of the reflex system and thereby minimize distortion. The effects of frequency bandwidth and amplitude of the displacement perturbations were tested in separate experiments. The coherence square, gain and phase between the EMG and angular displacement were calculated in order to characterize the stretch reflex under these conditions. It was found that the phase of the reflex response was dependent on both bandwidth and amplitude. For narrow-band displacements, the phase advance was about 30 degrees greater over the frequency range tested than for broad-band displacements, suggesting that the reflex response may be influenced by the predictability of the perturbation. At the smallest amplitude of 0.3 degrees, the peak phase advance was about 20 degrees greater than at the largest amplitude of 4.2 degrees. The gain was also higher and rose more steeply with frequency at smaller amplitudes. In the frequency range up to 12 Hz, the tonic stretch reflex responds most effectively to smaller-amplitude, more regular, higher-frequency inputs and this is consistent with a role for the reflex in counteracting small-amplitude oscillations, tremors and errors of voluntary movement.

Adult↗

Habituation and conditioning of the human long latency stretch reflex.

The effects of stretch repetition rate, prior warning stimuli and self administered stretch were examined on the size of the short and long latency components of the stretch reflex electromyographic EMG response in flexor pollicis longus and the flexor muscles of the wrist and fingers. Stretches of constant velocity and extent were given every 10 s, 5 s, 2 s, or 1 s to either the wrist or thumb during a small background contraction of the flexor muscles. The size of the long latency component of the stretch reflex (measured as the area under the averaged rectified EMG responses) declined dramatically at faster repetition rates, especially in the wrist and finger flexors. The size of the short latency component was relatively unaffected. The size of the electrically elicited H-reflex in forearm muscles also failed to habituate under the same conditions. If each individual trial of a series was examined, the long latency component of the stretch reflex EMG could be seen to decrease in size over the first three to six stretches if stretches were given every 1 s, but not if stretches were given every 10 s. When stretches were given every 5 s to either wrist or thumb, an electrical stimulus applied to the digital nerves of the opposite hand 1 s before stretch reduced the size of the long latency component of the reflex EMG response. The short latency component was unaffected.(ABSTRACT TRUNCATED AT 250 WORDS)

Adult↗

The influence of experimental muscle pain on the human soleus stretch reflex during sitting and walking.

OBJECTIVES: The stretch reflex is functionally important during human locomotion. Muscle pain has been found to increase the stretch reflex amplitude during sitting, possibly due to an altered fusimotor drive. To further study the importance of altered fusimotor activity due to muscle pain we investigated the combined effect of muscle pain and motor task on the soleus stretch reflex. METHODS: Stretch reflexes were elicited before, during and after experimentally induced muscle pain in soleus (i.m. infusion of 6% saline) in 3 experiments: (1) in the relaxed soleus muscle and before, during and after an isometric ramp contraction (500 ms, 0-10 Nm), (2) at 3 different time periods during walking, and (3) at matched pain intensity and soleus activity during sitting and walking. RESULTS: Infusion of hypertonic saline into the soleus muscle caused a significant facilitated stretch reflex in the relaxed muscle (P<0.01), but not during walking or during sitting and walking at matched soleus EMG and matched pain levels. The infusion of isotonic saline (non-painful) did not cause any changes (P = 0.75). CONCLUSIONS: The main findings of the present study were that experimental muscle pain facilitated the stretch reflex during pain in the relaxed muscle, but caused no changes in stretch reflex amplitude during sitting and walking at higher "functional" background EMG levels.

Adolescent↗