A study of synaptic connection between low threshold afferent fibres in common peroneal nerve and motoneurones in human tibialis anterior

We have induced H-reflex responses in human tibialis anterior motor units and analysed the results using the classical technique, peristimulus time histogram (PSTH), and a new technique, peristimulus frequencygram (PSF). The PSF has recently been shown to be more reliable than the PSTH for indicating the synaptic connections on motoneurones, and therefore we wished to examine the differences between the two analysis methods. Experiments were conducted on eleven healthy subjects (7 males and 4 females) who did not have any known neurological disorder. The subject sat comfortably on a dental chair and the common peroneal nerve was stimulated. In each experiment, about 600 electrical stimuli were applied to the nerve randomly between 1 and 2 s. The recordings were taken with both by surface electromyogram (SEMG) and as single motor unit potentials. We found that, when a stimulus induces an H-reflex, it also generates a period of reduced activity (silent period) and a long latency excitation in the PSTH. However, the PSF records in general do not match the indications of the PSTH records. For example, when the PSTH indicated existence of a silent period immediately following the H-reflex response, the discharge rate of the unit was in fact higher than the prestimulus rate. On the contrary, during the PSTH illustrated long latency excitatory response, the discharge rate was lower than the prestimulus rate. Our findings suggest that PSF gives significantly different results compared with the PSTH in determining the synaptic connection of the low threshold muscle afferents to the motoneurones. While PSTH indicated that there was a silent period immediately after the H-reflex, the PSF demonstrated that the silent period was actually a continuation of the net excitatory effect and not a genuine inhibition since the small number of action potentials occured during this period displayed higher discharge rates than the prestimulus level. Furthermore, the long latency excitation, as it was indicated in the PSTH; was actually a net inhibitory effect since the large number of spikes that occured during that period had lower discharge rates than the prestimulus average. In the lights of the recent brain slice findings and completely different results obtained using the two analysis techniques, we suggest that the PSF analysis should be used along with the PSTH to illustrate the net synaptic connection between peripheral receptors and motoneurones in the human nervous system.     

Effects of large excitatory and inhibitory inputs on motoneuron discharge rate and probability.

We elicited repetitive discharge in hypoglossal motoneurons recorded in slices of rat brain stem using a combination of a suprathreshold injected current step with superimposed noise to mimic the synaptic drive likely to occur during physiological activation. The effects of repetitive en mass stimulation of afferent nerves were simulated by the further addition of trains of injected current transients of varying shapes and sizes. The effects of a given current transient on motoneuron discharge timing and discharge rate were measured by calculating a peristimulus time histogram (PSTH) and a peristimulus frequencygram (PSF). The amplitude and time course of the simulated postsynaptic potentials (PSPs) produced by the current transients were calculated by convolving the current transient with an estimate of the passive impulse response of the motoneuron. We then compared the shape of the injected current transient and the simulated PSP to the profiles of the PSTH and the PSF records. The PSTHs produced by excitatory PSPs (EPSPs) were characterized by a large, short-latency increase in firing probability that lasted slightly longer than the rising phase of the EPSP, followed by a reduced discharge probability during the falling phase of the EPSP. In contrast, the PSF analysis revealed a proportionate increase in discharge rate over the entire profile of the EPSP, even though relatively few spikes occurred during the falling phase. The PSTHs associated with inhibitory PSPs (IPSPs) indicated a reduction in discharge probability during the initial, hyperpolarizing phase of the IPSP, followed by an increase in the discharge probability during its subsequent repolarizing phase. Using the PSF analysis, the initial phase of the IPSP appeared as a large hole in the record where a very small number or no discharges occurred. The subsequent phase of the IPSP was associated with frequency values that were lower than the background values. The primary features of both PSTHs and PSFs can be used to estimate the relative amplitudes of the underlying EPSPs and IPSPs. However, PSTHs contain secondary peaks and troughs that are not directly related to the underlying PSP but instead reflect the regular recurrence of spikes following those affected by the PSP. The PSF analysis is more useful for indicating the total duration and the profile of the underlying PSP. The shape of the underlying PSP can be obtained directly from the PSF records because the discharge frequency of the spikes follow the PSPs very closely, especially for EPSPs.     

Experimental skin pain and muscle pain induce distinct changes in human trigeminal motoneuronal excitability

Seeking information on the physiological properties of the trigeminal motoneuronal pool we investigated changes in the excitability of trigeminal motor system induced by two types of experimental pain (muscle and skin). In one session, we studied the effect of muscle pain induced by hypertonic saline infusion into the masseter muscle on the recovery cycle of the heteronymous H-reflex in the temporalis muscle and the homonymous silent period (SP) in the masseter muscle, both elicited by stimulation of the masseteric nerve in ten-healthy subjects. In another session, we studied the effect of laser stimuli applied to the perioral region, at conditioning intervals from 20 to 160 ms, on the temporalis H-reflex and masseter SP in nine healthy subjects. Whereas laser-induced skin pain significantly inhibited the temporalis H-reflex and facilitated the masseter SP (P < 0.01), muscle pain left the time course of the temporalis H-reflex and masseter SP unchanged (P > 0.05). The timing of temporalis H-reflex suppression and masseter-SP enhancement induced by laser stimuli indicates that facial skin nociceptors inhibit trigeminal motoneurones via multysynaptic reflex pathways. Hypertonic saline, a stimulus that predominantly activates group III and IV afferents, left both variables reflecting trigeminal motoneuron excitability unchanged. Due to the differences between the two experimental models, we cannot conclude that such inhibitory reflex pathway does not exist from muscle nociceptors to trigeminal motoneurones.     

Comparison of the inhibitory response to tendon and cutaneous afferent stimulation in the human lower limb

A powerful early inhibition is seen in triceps surae after transcutaneous electrical stimulation of the Achilles tendon [tendon electrical stimulation (TES)]. The aim of the present study was to confirm results from surface electromyogram (SEMG) recordings that the inhibition is not wholly or partly due to stimulation of cutaneous afferents that may lie within range of the tendon electrodes. Because of methodological limitations, SEMG does not reliably identify the time course of inhibitory and excitatory reflex components. This issue was revisited here with an analysis of changes in single motor unit (SMU) firing rate [peristimulus frequencygram (PSF)] and probability [peristimulus time histogram (PSTH)] to reexamine the time course of inhibitory SMU events that follow purely cutaneous (superficial sural) nerve stimulation. Results were then compared with similar data from TES. When compared with the reflex response to TES, sural nerve stimulation resulted in a longer onset latency of the primary inhibition and a weaker effect on SMU firing probability and rate. PSF also revealed that decreased SMU firing rates persisted during the excitation phase in SEMG, suggesting that the initial inhibition was more prolonged than previously reported. In a further study, the transcutaneous SEMG Achilles tendon response was compared with that from direct intratendon stimulation with insulated needle electrodes. This method should attenuate the SEMG response if it is wholly or partly dependent on cutaneous afferents. However, subcutaneous stimulation of the tendon produced similar components in the SEMG, confirming that cutaneous afferents made little or no contribution to the initial inhibition following TES.     

Ia reflexes and EPSPs in human soleus motor neurones

Summary The reflex responses of single motor units in the soleus muscle to electrical stimulation of the tibial nerve were recorded in human volunteers. A feature of the experiments was the stimulation paradigm used. In order to control the peristimulus firing rate, a computer triggered the stimulus isolator only when 2 interspike intervals of specified duration occurred in succession. In addition, the timing of the stimulus in relation to the preceding action potential was controlled in a manner similar to a conditioning/testing paradigm. The general pattern of response was an initial, H-reflex excitation at monosynaptic latency, followed first by a silent period due to the refractoriness of the motor neurone, then by other phases of reduced activity. When the stimulus intensity was increased, the intensity of the excitation and the duration of the silent period increased in parallel. When the pre-stimulus firing rate of the motor unit was varied, the amplitude of the H-reflex response, normalized to the number of stimulus trials, was similar at 6, 8 and 10 Hz, but was greater at 4 Hz in most units tested. These findings were consistent with a simple model of the events occurring at the cell membrane in this reflex which was proposed by Ashby and Zilm (1982a), although some modification of the model was necessary to account for the different response at 4 Hz. The improved stimulation paradigm enabled a direct estimate to be made of the amplitude and shape of the rising phase of the Ia EPSP in human motor neurones.     

Modulation, probably presynaptic in origin, of monosynaptic Ia excitation during human gait

Modulation of presynaptic inhibition of Ia afferents projecting monosynaptically to soleus motoneurones was investigated during human gait. Changes in presynaptic inhibition of Ia afferents were deduced from alterations in the amount of heteronymous soleus H-reflex facilitation evoked by a constant femoral nerve stimulation. It has been shown that this facilitation is mediated through a monosynaptic Ia pathway and that during its first 0.5 ms it is still uncontaminated by any polysynaptic effect and can be used to assess ongoing presynaptic inhibition of Ia terminals to soleus motoneurones. During gait, heteronymous facilitation was reduced with respect to its control value (rest during sitting) and modulated during the step cycle: it reached its maximum at mid-stance and decreased to near zero by the end of stance. At the same time the H-reflex amplitude was to some extent similarly modulated. It is argued that this decrease in heteronymous Ia facilitation and in H-reflex amplitude reflects an increased, ongoing presynaptic inhibition of Ia terminals projecting onto soleus motoneurones, which could be from central and/or peripheral origin. D1 inhibition, i.e. the late and long-lasting inhibition of the soleus H-reflex evoked by a train of stimuli to the common peroneal nerve, was used as another method to assess presynaptic inhibition. This D1 inhibition was decreased during gait, and it is argued that this decrease might reflect an occlusion in presynaptic pathways or increased presynaptic inhibition of pathways mediating the conditioning volley.     

Excitatory and inhibitory effects on human spinal motoneurones from magnetic brain stimulation

Stimulation of baboon motor cortex causes in the motoneurones (MNs) of intrinsic hand muscles monosynaptic excitatory postsynaptic potentials (EPSPs) and a disynaptic inhibitory postsynaptic potential (IPSP). These phenomena have been investigated in human MNs by applying pulsed magnetic stimuli over the scalp at random times during the tonic discharge of single hand muscle motor units (MUs). Post-stimulus time histograms (PSTHs) demonstrated an increased firing probability at between 25 and 35 ms. This major firing peak showed a multimodal form with interpeak intervals of 1.4-1.8 ms. When MUs were not fired by the stimulus, they were nevertheless inhibited from firing spontaneously. There are thus short latency excitatory and inhibitory cortical inputs to human MNs.     

On the mechanism of the post-activation depression of the H-reflex in human subjects

It was demonstrated that the soleus H-reflex was depressed for more than 10 s following a preceding passive dorsiflexion of the ankle joint. This depression was caused by activation of large-diameter afferents with receptors located in the leg muscles, as an ischaemic block of large-diameter fibres just below the knee joint abolished the depression, whereas a similar block just proximal to the ankle joint was ineffective. The depression of the H-reflex was not caused by changes in motoneuronal excitability, as motor-evoked potentials by magnetic brain stimulation were not depressed by the same passive dorsiflexion. Therefore it was concluded that the long-lasting depression is due to mechanisms acting at presynaptic level. The transmission of the monosynaptic Ia excitation from the femoral nerve to soleus motoneurones was not depressed by the ankle dorsiflexion. The depression thus seems to be confined to those afferents that were activated by the conditioning dorsiflexion. In parallel experiments on decerebrate cats, more invasive methods have complemented the indirect techniques used in the experiments on human subjects. A similar long-lasting depression of triceps surae monosynaptic reflexes was evoked by a preceding conditioning stimulation of the triceps surae Ia afferents. This depression was accompanied by a reduction of the monosynaptic Ia excitatory postsynaptic potential recorded intracellularly in triceps surae motoneurones, but not by changes in the input resistance or membrane potential in the motoneurones. Stimulation of separate branches within the triceps surae nerve demonstrated that the depression is confined to those afferents that were activated by the conditioning stimulus. This long-lasting depression was not accompanied by a dorsal root potential. It is concluded that the long-lasting depression is probably caused by a presynaptic effect, but different from the classical GABAergic presynaptic inhibition which is widely distributed among afferent fibres and accompanied by dorsal root potentials. It is more probably related to the phenomenon of a reduced transmitter release from previously activated fibres, i.e. a homosynaptic post-activation depression. The consequences of this post-activation depression for the interpretation of results on spinal mechanisms during voluntary movements in man are discussed.     

Monosynaptic and oligosynaptic contributions to human ankle jerk and H-reflex

Studies were undertaken in normal subjects to determine whether it is possible for oligosynaptic reflex pathways to affect motoneuron discharge in the ankle jerk and H-reflex of the soleus. It is argued that if the rising phase of the increase in excitability of the soleus motoneuron pool produced by tendon percussion or by electrical stimulation of the peripheral nerve lasts more than a few milliseconds and if the increase in excitability takes several milliseconds to reach the threshold for motoneuron discharge, these reflexes are unlikely to be exclusively monosynaptic. In relaxed subjects, changes in excitability of the soleus motoneuron pool produced by tendon percussion and by electrical stimulation of the tibial nerve were examined using conditioning stimuli just below threshold and a test H-reflex just above threshold for a reflex response. The increase in excitability due to tendon percussion had an average rise time of 10.8 ms and a total duration of approximately 25 ms. With electrical stimulation the rising phase appeared shorter, but it could not be measured accurately due to afferent refractoriness. In single motor units, the rise times of the composite excitatory postsynaptic potentials (EPSPs) set up by subthreshold tendon percussion and by subthreshold electrical stimulation of the tibial nerve were estimated from changes in the probability of discharge of voluntarily activated single motor units. Rise times were significantly longer with tendon percussion (mean +/- SD, 7.1 +/- 2.3 ms; n = 34) than with electrical stimulation (2.4 +/- 1.4 ms; n = 32). In four experiments in which a number of motor units were studied using identical mechanical and identical electrical stimuli, the poststimulus time histograms (PSTHs) for each stimulus were pooled to provide an estimate of the rise time of the excitability change in the motoneuron pool. The mean rise times of these four samples were 10.5 ms with mechanical stimulation and 4.5 ms with electrical stimulation. The spontaneous variability in latency of reflexly activated single motor units was 0.8-3.1 ms (average SD, 0.34 ms) in the tendon jerk, and 0.6-1.4 ms (average SD, 0.19 ms) in the H-reflex. Comparison of these figures with the measurements of rise time given above suggests that the composite EPSPs are larger than the background synaptic noise. With six motor units, the timing of reflex discharge in the tendon jerk when the subject was relaxed was compared with the timing of the change in probability of discharge due to apparently identical percussion when the units were activated voluntarily.(ABSTRACT TRUNCATED AT 400 WORDS)     

Long-loop reflex from arm afferents to remote muscles in normal man

The present study was designed to examine the effects of median nerve stimulation on motoneurones of remote muscles in healthy subjects using H-reflex, averaged EMG and PSTH methods. Stimulation of the median nerve induced facilitation of soleus H-reflex from about 50 ms and it reached a peak at about 100 ms of conditioning-test interval. Afferents that induced the facilitation consisted of at least two types of fibres, the high-threshold cutaneous fibres and the low-threshold fibres. When the effects were examined by the averaged surface EMG and PSTH, no facilitation but rather inhibition or inhibition-facilitation was induced in all tested muscles except for the upper limb muscles on the stimulated side. The inhibition latency was shortest in masseter muscle and longest in leg muscles, while values for the contralateral upper limb muscles were in the middle, indicating that the onset of inhibition was delayed from rostral to caudal muscles. Inputs from the median nerve converged to inhibitory interneurones, which mediate the masseter inhibitory reflex. Our findings suggested that inputs from the median nerve initially ascend to the brain, at least to the brainstem, and then descend to the spinal cord. Therefore, inhibition induced by median nerve stimulation was not considered as an interlimb reflex mediated by a propriospinal pathway, but long-loop reflex, at least via the pons. The discrepancy between the results of reflex and motor units suggests that facilitation of soleus H-reflex following median nerve stimulation was mainly due to reduced presynaptic inhibition.     

Modulation of motor unit discharge rate and H-reflex amplitude during submaximal fatigue of the human soleus muscle

Declining motor unit discharge rates and H-reflex amplitude have been observed in separate experiments during fatiguing submaximal contractions in humans. The purpose of this experiment was to investigate motor unit discharge rate, H-reflex amplitude, and twitch contractile properties concurrently during a fatiguing submaximal isometric contraction of the ankle plantarflexors. Eleven healthy subjects performed fatiguing contractions of low force (25% maximal voluntary contraction (MVC)) or high force (42–66% MVC). Hoffmann (H)-reflexes, muscle compound action potentials (M-waves), twitch contractile properties, and motor unit discharges were recorded from the soleus muscle. In the low-force fatigue task, motor unit firing rate increased gradually over time, whereas the resting H-reflex was significantly depressed at 15% of endurance time and remained quasiconstant for the rest of the task. This suggests that the processes mediating the resting H-reflex depression are relatively independent of those modulating the motor unit firing rate during a low-force fatigue task. In the high-force fatigue task, a decline in the average motor unit discharge rate was accompanied by a decrease in the resting H-reflex amplitude and a prolongation of the twitch half-relaxation time (HRT) at the completion of the fatigue task. Overall, motor unit firing rate was modulated in parallel with changes in the twitch HRT, consistent with the muscle wisdom hypothesis.     

Reflex response and control of the human soleus and gastrocnemius muscles during walking and running at increasing velocity

We measured the soleus and the gastrocnemius H-reflex modulation in seven subjects during walking at 4.5 km/h and during running at 8, 12 and 15 km/h. The recordings in the medial gastrocnemius were corrected for cross-talk from the soleus muscle. The gastrocnemius H-reflex was in general lower than the soleus H-reflex. In both muscles the H-reflex increased significantly from walking to running but also with increasing running speed. The peak of EMG activity increased in both muscles with increasing speed. The V-wave of both muscles was absent or rather low during walking, but it increased significantly from walking to running with increasing running speed in the soleus but not in the medial gastrocnemius. In both muscles the V-wave was highest just prior to heel strike. It is suggested that this was due to a high firing frequency of the motoneurones in this phase of the movement. It is concluded that a shift towards the faster gastrocnemius at higher running speeds on behalf of the soleus muscle did not occur. The fact that the physiological cross-sectional area of the soleus is much larger than that of the lumped gastrocnemii is most probably the reason why the soleus is important also at higher running velocities.     

Synaptic connections from large muscle afferents to motoneurones in human lower limbs

OBJECTIVES: Our study was undertaken to investigate reciprocal inhibition in humans both from ankle flexors to extensors and from ankle extensors to flexors. METHODES: Changes in the firing probability of single motor units in response to electrical stimulation of muscle nerves (the peristimulus time histogram technique) were used to derive the reciprocal projections of muscle spindle Ia afferents to the motoneurones of ankle muscles. Discharges of units in ankle flexors (the tibialis anterior muscle [TA]) and extensors (soleus [SOL] and medial gastrocnemius [MG] muscles) were investigated respectively after stimulation of the posterior tibial (PTN) and common peroneal (CPN) nerves (predominantly on the deep branch). In eight normal subjects aged 24 to 40 years, one motor unit per each muscle was studied. RESULTS: CPN stimulation produced reciprocal Ia inhibition in the SOL of 5 of 7 of them and in the MG of 3 of 5, whereas PTN stimulation produced reciprocal Ia inhibition in the TA of only 2 of 6 subjects. CONCLUSIONS: These findings suggest that at low level contraction reciprocal Ia inhibition from ankle flexors to extensors may be stronger than that from ankle extensors to flexors.     

Reciprocal inhibition of soleus motor output in humans during walking and voluntary tonic activity

1. The extent to which an active, human motoneuron pool can be inhibited via short-latency inhibitory pathways was studied by stimulating the common peroneal nerve and recording the inhibition of on-going soleus electromyographic (EMG) activity. The responses were compared at the same EMG level during walking and tonic voluntary activity to determine whether the inhibition was task dependent. 2. In both tasks the amount of inhibition (measured as the depression in rectified, filtered, and averaged EMG activity) increased approximately linearly with the amount of motor activity, as determined from the mean EMG level before stimulation (correlation coefficient greater than or equal to 0.9). No difference in the amount of inhibition was found between the two tasks at the same stimulus and EMG levels. 3. Previously published studies based on the H-reflex method have reported that the amount of inhibition decreases with the amount of motor activity. On the contrary, single-unit studies and the present results suggest that segmental inhibitory reflexes retain their capacity to mediate a rapid reduction of motoneuronal discharge during voluntary activity. This inhibition may be important in regulating the amount of activity early in the stance phase of walking and during the transition from stance to the swing phase. 4. Analytic results are derived in an APPENDIX that should be of general interest in interpreting the inhibition of motor units from a peristimulus time histogram (PSTH). The linear correlation between inhibition and level of voluntary activity can be explained if newly recruited units are strongly inhibited by the stimulus, whereas previously active motor units are inhibited relatively less, as their firing rate increases with increasing background activity.     

Phase-response curves give the responses of neurons to transient inputs

Neuronal firing is determined largely by incoming barrages of excitatory postsynaptic potentials (EPSPs), each of which produce a transient increase in firing probability. To measure the effects of weak transient inputs on firing probability of cortical neurons, we compute phase-response curves (PRCs). PRCs, whose shape can be related to the dynamics of spike generation, document the changes in timing of spikes caused by an EPSP in a repetitively firing neuron as a function of when it arrives in the interspike interval (ISI). The PRC can be exactly related to the poststimulus time histogram (PSTH) so that knowledge of one uniquely determines the other. Typically, PRCs have zero values at the start and end of the ISI, where EPSPs have minimal effects and a peak in the middle. Where the peak occurs depends in part on the firing properties of neurons. The PRC can have regions of positivity and negativity corresponding respectively to speeding up and slowing down the time of the next spike. A simple canonical model for spike generation is introduced that shows how both the background firing rate and the degree of postspike afterhyperpolarization contribute to the shape of the PRC and thus to the PSTH. PRCs in strongly adapting neurons are highly skewed to the right (indicating a higher change in probability when the EPSPs appear late in the ISI) and can have negative regions (indicating a decrease in firing probability) early in the ISI. The PRC becomes more skewed to the right as the firing rate decreases. Thus at low firing rates, the spikes are triggered preferentially by inputs that occur only during a small time interval late in the ISI. This implies that the neuron is more of a coincidence detector at low firing frequencies and more of an integrator at high frequencies. The steady-state theory is shown to also hold for slowly varying inputs.     

The relative dependence of the activity of renshaw cells on recurrent pathways during contraction of the triceps muscle

Renshaw cell activity was recorded simultaneously with motoneuronal unit discharge during vibration and tetanic stimulation of triceps muscles in decerebrated cats. The experiments confirm that, in this preparation, the motoneurones are the main source of Renshaw cell firing during muscle stretch and vibration and when motoneuronal discharge was induced through the gamma loop. However they also show that a discharge of Renshaw cells, monosynaptically coupled with triceps motoneurones through their recurrent collaterals, could be elicited during contraction of the muscle at the time when the discharge of these motoneurones had been silenced.The recording of the stretch receptors and motoneuronal unit discharge during stretch, vibration, and ventral root stimulation gave evidence of the contribution of the withdrawal of excitation by primary endings to the occurrence of the silent period during tetanic contraction of the muscle. The measurements of the critical firing level in motoneuronal units responding reflexly to held stretch and vibration of the muscles, and silencing their discharge during muscle shortening, showed that these cells are amongst the lowest ranking in the pool. For these reasons, these data suggest that Renshaw cell firing during vibration and tetanic contraction of the muscle cannot be attributed only to the alpha motoneurone excitation by the Ia fibres.     

Mechanisms regulating the activity of facial nucleus motoneurones--2. Synaptic activation from the caudal trigeminal nucleus

Field and postsynaptic potentials of facial motoneurones evoked by stimulation of the caudal trigeminal nucleus were studied in cats by means of extra- and intracellular recording. Mono- and polysynaptic input onto facial motoneurones from the caudal trigeminal nucleus were shown. Four types of responses were distinguished: excitatory postsynaptic potentials generating a single action potential; a gradual shift of depolarization inducing multiple discharges; a rhythmic discharge of action potentials appearing at a low level of depolarization; excitatory postsynaptic potentials or a sequence of excitatory and inhibitory postsynaptic potentials. Multiple discharge was shown to appear as a result of effective summation of high frequency excitatory influences from efferent neurones of the caudal trigeminal nucleus projecting into the facial nucleus. Factors facilitating the development of gradual depolarization are: dendritic localization of synaptic terminals, dendritic origin of after-depolarizing processes and the high input resistance of the facial motoneurone membrane. It is thought that specific features of facial motoneurones and properties of afferent inputs are supposed to provide high sensitivity of neuronal organization of the facial nucleus to afferent signals as well as wide diversity in controlling its activity.     

Responses of neurones in cat's abducens nuclei to horizontal angular acceleration

Summary Single cell activity was recorded extracellularly from neurones in cat's abducens nuclei in unanesthetized, decerebrate preparations. Two types of neurones were found. i) The motoneurones (type II_A) increased their firing linearly or exponentially to contralateral constant angular accelerations and decreased firing to ipsilateral rotation. The increased frequency was maintained at a constant level without any sign of adaptation. This constant level of maximum frequency was proportional to the logarithm of the acceleration applied. No significant differences of the frequency responses of tonic and silent motoneurones were found. The conduction velocities of the axons of spontaneously firing type II_A cells were slower than in silent neurones. Electrical stimulation of the contralateral VIIIth nerve led to activation of the type II_A cells; stimulation of the ipsilateral nerve resulted in inhibition of both spontaneous discharge and antidromic invasion of the motoneurones. ii) The type I_A units increased their firing rate to ipsilateral rotation and ceased to discharge in response to contralateral rotation. They never were excited antidromically by stimulation of the VIth nerve. However, they were excited by stimulation of the ipsilateral VIIIth nerve. Their possible functional role in the vestibulo-ocular system was discussed.Parts of this study are subject of a dissertation to be presented to the Naturwissenschaftliche Fakultät, Universität Frankfurt.     

Analysis of motoneuron responses to composite synaptic volleys (computer simulation study)

This paper deals with the analysis of changes in motoneuron (MN) firing evoked by repetitively applied stimuli aimed toward extracting information about the underlying synaptic volleys. Spike trains were obtained from computer simulations based on a threshold-crossing model of tonically firing MN, subjected to stimulation producing postsynaptic potentials (PSPs) of various parameters. These trains were analyzed as experimental results, using the output measures that were previously shown to be most effective for this purpose: peristimulus time histogram, raster plot and peristimulus time intervalgram. The analysis started from the effects of single excitatory and inhibitory PSPs (EPSPs and IPSPs). The conclusions drawn from this analysis allowed the explanation of the results of more complex synaptic volleys, i.e., combinations of EPSPs and IPSPs, and the formulation of directions for decoding the results of human neurophysiological experiments in which the responses of tonically firing MNs to nerve stimulation are analyzed.     

Evidence for excitation of the human lower limb motoneurones by group II muscle afferents

The possibility was investigated that stimulation of high-threshold afferents in the common peroneal nerve (CPN) evokes excitation of quadriceps (Q) motoneurones in humans. Effects of conditioning stimuli at motor threshold (1×MT) and at higher intensities were compared on both the Q H-reflex and the post-stimulus time histogram (PSTH) of individual motor units. At 1×MT, CPN stimulation evokes a facilitation, which has been shown to be caused by an interneuronally mediated group I excitation. Increasing the CPN stimulus intensity above 2×MT caused this early excitation to increase and a later facilitation to appear both in the H-reflex and in the PSTH of single units. The later excitation had its threshold between 2 and 3×MT, and it appeared 4–8 ms after the group I-induced excitation. The higher threshold and the longer latency suggest that this excitation is evoked by afferents with a smaller diameter than group I afferents, and group II afferents meet this criterion.