C. Crone

Sensitivity of monosynaptic test reflexes to facilitation and inhibition as a function of the test reflex size: a study in man and the cat

SummaryIn parallel experiments on humans and in the cat it was investigated how the sensitivity of monosynaptic test reflexes to facilitation and inhibition varies as a function of the size of the control test reflex itself. In man the monosynaptic reflex (the Hoffmann reflex) was evoked in either the soleus muscle (by stimulation of the tibial nerve) or the quadriceps muscle (by stimulation of the femoral nerve). In the decerebrate cat monosynaptic reflexes were recorded from the nerves to soleus and medial gastrocnemius muscles; they were evoked by stimulation of the proximal ends of the sectioned L7 and S1 dorsal roots. Various excitatory and inhibitory spinal reflex pathways were used for conditioning the test reflexes (e.g. monosynaptic Ia excitation, disynaptic reciprocal inhibition, cutaneous inhibition, recurrent inhibition, presynaptic inhibition of the Ia fibres mediating the test reflex). It was shown that the additional number of motoneurones recruited in a monosynaptic test reflex by a constant excitatory conditioning stimulus was very much dependent on the size of the test reflex itself. This dependency had the same characteristic pattern whatever the conditioning stimulus. With increasing size of the test reflex the number of additionally recruited motoneurones first increased, then reached a peak (or plateau) and finally decreased. A similar relation was also seen with inhibitory conditioning stimuli. The basic physiological factors responsible for these findings are discussed. Finally, the implications for the interpretation of experiments in man with the H-reflex technique are considered.

Reciprocal Ia inhibition between ankle flexors and extensors in man.

1. Reciprocal inhibition between antagonist muscle groups at the ankle has been investigated in sixty healthy subjects. Hoffmann reflexes (H reflexes) in the soleus and tibialis anterior muscles were used to assess changes in reciprocal inhibition evoked by electrical stimulation of antagonist muscle nerves. 2. Inhibition of the soleus H reflex was evoked by a single conditioning stimulus to the common peroneal nerve, and inhibition of the tibialis anterior H reflex was elicited by one conditioning stimulus to the posterior tibial nerve. Symmetrical central connections between the antagonist flexors and extensors were assumed and under this assumption the central delay for the inhibition, in addition to the delay for monosynaptic Ia excitation, was calculated to be about 1 ms. The inhibition was evoked by weak stimuli to the nerves from antagonist muscle groups; the threshold for the inhibition was around 0.6 X threshold for a direct motor response (M‐threshold). Furthermore, tendon taps to the Achilles tendon facilitated the soleus H reflex and inhibited the tibialis anterior reflex at short latencies. The short central delay, the low electrical threshold and the.actions of Achilles tendon taps strongly suggest that the early reciprocal inhibition is homologous to the disynaptic Ia inhibition previously studied in animal experiments. 3. With the test soleus H reflex kept at 15‐25% of the maximum directly evoked motor response (M‐response) and the strength of the conditioning peroneal nerve stimulation kept at 1.0 X M‐threshold, the inhibition from the peroneal nerve ranged between 0 and 40% (mean, 14.9%) at rest. 4. Changes in the amount of reciprocal inhibition from the peroneal nerve were studied both during tonic and dynamic dorsi‐ and plantarflexion. During tonic dorsiflexion there was no significant change of inhibition as compared to rest, while inhibition decreased during tonic plantarflexion. However, during ramp‐and‐hold dorsiflexion there was a transient increase in reciprocal inhibition of the soleus H reflex. This increase in inhibition from the peroneal nerve could be seen 50 ms prior to the onset of contraction. The increase in inhibition before and at the very beginning of the contraction cannot be due to sensory feed‐back during contraction, but must depend on a supraspinal control of the spinal cord. 5. At conditioning‐test intervals of 4‐6 ms, the inhibition of the soleus H reflex from the peroneal nerve was considerably larger during tonic dorsiflexion than at rest. Thus, tonic dorsiflexion revealed an inhibition with long latency from the peroneal nerve, which was not seen at rest.(ABSTRACT TRUNCATED AT 400 WORDS)

The spinal pathophysiology of spasticity - from a basic science point of view

Spasticity is a term, which was introduced to describe the velocity‐sensitive increased resistance of a limb to manipulation in subjects with lesions of descending motor pathways. This distinguishes spasticity from the changes in passive muscle properties, which are often seen in these patients, but are not velocity‐sensitive. Increased excitability of the stretch reflex is thus a central component of the definition of spasticity. This review describes changes in cellular properties and transmission in a number of spinal reflex pathways, which may explain the increased stretch reflex excitability. The review focuses mainly on results derived from the use of non‐invasive electrophysiological techniques, which have been developed during the past 20–30 years to investigate spinal neuronal networks in human subjects, but work from animal models is also considered. The reflex hyperexcitability develops over several months following the primary lesion and involves adaptation in the spinal neuronal circuitries caudal to the lesion. In animal models, changes in cellular properties (such as ‘plateau potentials’) have been reported, but the relevance of these changes to human spasticity has not been clarified. In humans, numerous studies have suggested that reduction of spinal inhibitory mechanisms (in particular that of disynaptic reciprocal inhibition) is involved. The inter‐subject variability of these mechanisms and the lack of objective quantitative measures of spasticity have impeded disclosure of a clear causal relationship between the alterations in the inhibitory mechanisms and the stretch reflex hyperexcitability. Techniques which make such a quantitative measure possible as well as longitudinal studies where development of reflex excitability and changes in the inhibitory mechanisms are followed over time are in great demand.

Methodological implications of the post activation depression of the soleus H-reflex in man

SummaryA long lasting inhibition (> 8 s) of the soleus Hoffmann reflex (H-reflex) was evoked by a preceding soleus H-reflex, by a brief voluntary ankle flexor or extensor muscle contraction or by a tap applied to the Achilles tendon. The time course of this long lasting inhibition was similar in all these cases, suggesting that the same spinal mechanism is involved. Furthermore, it was shown that the post-activation depression may interfere with the determination of inhibitory or facilitatory effects on the H-reflex. It is stressed that when the onset of inhibitory or facilitatory effects on the soleus H-reflex is to be determined in relation to start of an ankle movement, either very long stimulus intervals (> 8 s) must be used, or the onset must be determined in relation to a reference value of the soleus H-reflex, which may be influenced by the long lasting inhibitory effect, but not yet by the succeeding muscle contraction.

Maintained changes in motoneuronal excitability by short‐lasting synaptic inputs in the decerebrate cat.

1. During investigation of the tonic stretch reflex in the unanaesthetized decerebrate cat we observed that a short train of impulses in Ia afferents from the soleus muscle (or its synergists) may cause a prolonged activity in the soleus muscle as judged by EMG and tension recordings. This excitability increase, which outlasted the stimulus train, could stay virtually constant during long periods (even minutes), but could be terminated at any time by a train of impulses in, for example, the peroneal nerve. 2. Gradation of the strength of stimulation and the duration of the train of impulses show that the amount of maintained excitability increase depends‐within some limits‐on the total amount of Ia impulses. 3. In paralysed preparations a short train of impulses in Ia afferents from any part of the triceps surae, caused a maintained increase of the efferent activity in the nerves to triceps surae and a maintained increase of the triceps surae monosynaptic test reflex. These experiments demonstrate the existence of a central mechanism (in the spinal cord and/or the brain stem), which is responsible for the maintained excitability increase seen in motoneurones to the homonymous and synergic muscles. 4. In acute spinal preparations it was not possible to demonstrate any long‐lasting excitability increase by a train of Ia impulses. Following intravenous administration of the serotonin precursor 5‐hydroxytryptophan, mimicking the tonic activity of these pathways in the decerebrate state, it was again possible to elicit the long‐lasting excitability increase by a train of impulses in Ia afferents. A subsequent I.V. injection of methysergide (a serotonin receptor blocker) abolished the long‐lasting excitability increase. This set of experiments demonstrates that the basic mechanism responsible for the maintained excitability increase is located at segmental level, and involves serotonergic systems. 5. It was demonstrated that activation of several ipsilateral and crossed reflex pathways by trains of impulses in cutaneous or high‐threshold muscle afferents could trigger a maintained excitability increase of those motoneurone pools which were activated by the stimulation. Trains of stimuli to facilitatory regions in the brain stem could also cause a long‐lasting excitability increase of motoneurones. Furthermore, activation of all reflex pathways which mediate postsynaptic inhibition to a motor nucleus (including recurrent inhibition via Renshaw cells) could terminate the prolonged excitability increase of that particular motor nucleus.(ABSTRACT TRUNCATED AT 400 WORDS)

H-reflexes are smaller in dancers from The Royal Danish Ballet than in well-trained athletes

SummaryThe size of the maximalH-reflex (Hmax) was measured at rest and expressed as a percentage of the maximalM-response (Mmax) in 17 untrained subjects, 27 moderately trained subjects, 19 well-trained subjects and 7 dancers from the Royal Danish Ballet. TheHmax/Mmax was significantly larger in the moderately and well-trained subjects than in the untrained subjects but smaller in the ballet dancers. It is therefore suggested that both the amount and the type of habitual activity may influence the excitability of spinal reflexes.

Spinal mechanisms in man contributing to reciprocal inhibition during voluntary dorsiflexion of the foot.

1. The inhibition of the soleus Hoffmann reflex (H reflex) during voluntary dorsiflexion of the foot‐‐henceforth referred to as ‘natural’ reciprocal inhibition‐‐was found to be initiated 50 ms before the onset of the EMG activity in the tibialis anterior muscle and to increase gradually during a ramp‐and‐hold dorsiflexion. There was a positive correlation between strength of tonic dorsiflexion and amount of ‘natural’ reciprocal inhibition. 2. The change of activity in the disynaptic and a long‐latency group Ia inhibitory pathway and the change in presynaptic inhibition of the Ia fibres mediating the soleus H reflex were tested separately during ramp‐and‐hold dorsiflexion as well as during tonic dorsiflexion of the foot, and the results were compared with the development of the ‘natural’ reciprocal inhibition of the unconditioned soleus H reflex. 3. The disynaptic group I inhibition of soleus motoneurones was increased, as compared to rest, during the dynamic phase of a ramp‐and‐hold dorsiflexion movement, but the inhibition generally did not increase during tonic dorsiflexion of the foot. 4. The long‐latency group I inhibition was seen only during dorsiflexion of the foot. It appeared around 50 ms before tibial anterior EMG activity and there was a positive correlation between strength of tonic dorsiflexion and amount of this long‐latency inhibition. 5. Presynaptic inhibition of Ia afferents terminating on soleus motoneurones was estimated by an indirect method. The increase of presynaptic inhibition started soon after the onset of the ramp‐and‐hold dorsiflexion, and gradually became more pronounced during the ramp phase. The amount of presynaptic inhibition was positively correlated with strength of tonic dorsiflexion. 6. It is concluded that all investigated mechanisms may contribute to the ‘natural’ reciprocal inhibition and it seems that the different pathways are used differentially during different types of movement.

Post-activation depression of Soleus stretch reflexes in healthy and spastic humans

Reduced depression of transmitter release from Ia afferents following previous activation (post-activation depression) has been suggested to be involved in the pathophysiology of spasticity. However, the effect of this mechanism on the myotatic reflex and its possible contribution to increased reflex excitability in spastic participants has not been tested. To investigate these effects, we examined post-activation depression in Soleus H-reflex responses and in mechanically evoked Soleus stretch reflex responses. Stretch reflex responses were evoked with consecutive dorsiflexion perturbations delivered at different intervals. The magnitude of the stretch reflex and ankle torque response was assessed as a function of the time between perturbations. Soleus stretch reflexes were evoked with constant velocity (175°/s) and amplitude (6°) plantar flexion perturbations. Soleus H-reflexes were evoked by electrical stimulation of the tibial nerve in the popliteal fossa. The stretch reflex and H-reflex responses of 30 spastic participants (with multiple sclerosis or spinal cord injury) were compared with those of 15 healthy participants. In the healthy participants, the magnitude of the soleus stretch reflex and H-reflex decreased as the interval between the stimulus/perturbation was decreased. Similarly, the stretch-evoked torque decreased. In the spastic participants, the post-activation depression of both reflexes and the stretch-evoked torque was significantly smaller than in healthy participants. These findings demonstrate that post-activation depression is an important factor in the evaluation of stretch reflex excitability and muscle stiffness in spasticity, and they strengthen the hypothesis that reduced post-activation depression plays a role in the pathophysiology of spasticity.

Amplitude of the maximum motor response (Mmax) in human muscles typically decreases during the course of an experiment

Abstract It was shown that the amplitude of the soleus Mmax and Hmax responses decreases in the course of long-lasting H-reflex studies. The peak-to-peak amplitudes of the Mmax and Hmax responses in the soleus muscle (and the Mmax in the tibialis anterior muscle and small hand muscles) were measured repeatedly for 1–3 h in 20 subjects. 3–5 Mmax responses and 5–10 Hmax responses were elicited about every 3 min while the subject was at rest. Decreases in the soleus Mmax response of up to 50.5% (mean 20.5% SEM 2.2) and of the soleus Hmax of up to 49.7% (mean 19.1% SEM 3.7) in relation to the amplitudes measured at the beginning of the experiment were seen in 17 subjects. In 3 subjects no Mmax amplitude decrease was seen. The maximum decrease was reached between 10 and 100 min (mean 44.2 min SEM 4.3). An Mmax amplitude decrease was also seen in the tibialis anterior muscle and in two small hand muscles. In some subjects the decrease of the Mmax response seemed to be initiated by the infrequent supramaximal stimulations. The possible causes for this amplitude reduction, as well as the methodological consequences of these findings for H-reflex studies and fatigue studies, are briefly discussed.

Reciprocal Ia inhibition from the peroneal nerve to soleus motoneurones with special reference to the size of the test reflex

SummaryThe aim of the study was to examine the supraspinal control during voluntary movements of the foot in man, of the Ia inhibitory interneurones activated from the anterior tibial muscle and projecting to the soleus α-motoneurones. Previous studies have reported an increased inhibition of the soleus α-motoneurones by a constant conditioning stimulus to the common peroneal nerve during dorsiflexion of the foot. This was interpreted as a sign of supraspinal facilitation of the Ia inhibitory interneurones. However, these results could not be reproduced in the present study. The contradictory results can probably be explained by some important methodological differences in the use of the H-reflex technique.