Spike-triggered Averaging Technique Research Articles

Ascending projections of posterior canal-activated excitatory and inhibitory secondary vestibular neurons to the mesodiencephalon in cats.

The axonal projections of 62 posterior canal (PC)-activated excitatory and inhibitory secondary vestibular neurons were studied electrophysiologically in cats. PC-related neurons were identified by monosynaptic activation elicited by electrical stimulation of the vestibular nerve and activation following nose-up rotation of the animal's head. Single excitatory and inhibitory neurons were identified by antidromic activation following electrical stimulation of the contralateral and ipsilateral medial longitudinal fasciculus, respectively. The oculomotor projections of identified neurons were confirmed with a spike-triggered averaging technique. The axonal projections of the identified neurons were then studied by systematic, antidromic stimulation of the mesodiencephalon. Excitatory neurons showed two main types of axonal projections. In one type, axonal branches were issued to the interstitial nucleus of Cajal, central gray, and thalamus including the ventral posterolateral, ventral posteromedial, ventral lateral, ventral medial, centromedian, central lateral, lateral posterior, and ventral lateral geniculate nuclei. The other type was more frequently observed, giving off axon collaterals to the above-mentioned regions and to Forel's field H as well. Inhibitory neurons issued axonal branches to limited areas which included the central gray, interstitial nucleus of Cajal, its adjacent reticular formation and caudalmost part of Forel's field H, but not the rostral part of the Forel's field H and the thalamus. These results suggest that PC-related excitatory neurons participate in the genesis of vertical eye movements and in the perception of the vestibular sensation, and that PC-related inhibitory neurons seem to take part only in the genesis of vertical eye movements.

Supra- and juxtatrigeminal inhibitory premotor neurons with bifurcating axons projecting to masseter motoneurons on both sides

Inhibitory neurons participating in the bilateral disynaptic inhibition of jaw-closing motoneurons by stimulation of unilateral trigeminal sensory branches were searched for in the reticular formation around the trigeminal motor nucleus in cats anaesthetized with pentobarbital. Extracellular recordings were made from neurons which responded orthodromically after a monosynaptic latency to single shock stimulation of the ipsilateral infraorbital and/or inferior alveolar nerves. Direct inhibitory connection with contralateral masseter motoneurons was demonstrated in reticular neurons by the spike-triggered averaging technique, i.e., by averaging the intracellular potentials of a contralateral masseter motoneuron with respect to spontaneously occurring spikes of a reticular interneuron. By intraaxonal injection of neurobiotin, electrophysiologically identified inhibitory premotor reticular neurons were found to project to and to terminate in the trigeminal motor nuclei on both sides. Termination in the contralateral motor nucleus was demonstrated for four neurons that showed the peripheral input pattern stated above. The results provide hard evidence for contralaterally projecting interneurons in the reticular formation, participating in peripherally evoked disynaptic inhibition of masseter motoneurons on the contralateral side. Given the previously reported findings that the supratrigeminal region contains neurons which project to the ipsilateral motor nucleus and mediate disynaptic inhibition of masseter motoneurons, it is suggested that the supratrigeminal region contains bilaterally projecting interneurons, mediating peripherally evoked disynaptic inhibition of masseter motoneurons on both sides.

Motor unit estimation: Reproducibility of the spike‐triggered averaging technique in normal and ALS subjects

Reproducibility of the spike-triggered averaging technique of motor unit estimation (MUE) was assessed in biceps-brachialis muscle in 10 normal subjects and 15 subjects with amyotrophic lateral sclerosis (ALS). MUE was calculated by dividing the compound muscle action potential by the mean amplitude of 15 surface motor unit potentials (S-MUPs) of low recruitment threshold. Averaged MUE values in normal subjects were higher than in ALS subjects, with few values overlapping. Differences between test and retest MUE values were not significant for either subject group. The relative differences between test-retest values were 45.3% for normal subjects and 32.6% for ALS subjects. Correlation coefficients between test and retest values were low (r = 0.07) for normal subjects when influential outlying points were removed, and higher (r = 0.65) for ALS subjects when individuals with MUE values within the normal range were removed. The higher correlation of test-retest MUE values in ALS subjects compared to normal subjects may be due to a greater probability of resampling among the smaller number of motor units in ALS subjects. In summary, the reproducibility and technical aspects of the spike-triggered averaging technique are similar to those reported for other MUE techniques.

A physiological study of vestibular and prepositus hypoglossi neurones projecting to the abducens nucleus in the alert cat.

1. Vestibular and prepositus hypoglossi (PH) neurones projecting to the abducens (ABD) nucleus were recorded in the alert cat. Their discharge characteristics were analysed to ascertain the origin of the horizontal eye position signal present in ABD neurones. 2. Neurones were classified according to: their location with respect to the ABD nucleus; their antidromic activation from the ABD nucleus; the synaptic field potential they induced in the ABD nucleus with the spike-triggered averaging technique; and their activity during spontaneous and vestibularly induced eye movements. 3. Vestibular neurones projecting to the ABD nucleus were located in the rostral medial vestibular nucleus. They were excitatory on the contralateral and inhibitory on the ipsilateral ABD neurones. Both types of premotor vestibular neurone showed a firing rate weakly related to eye position, increasing for eye fixations in the contralateral on-direction, and decreasing with ipsilateral fixation. Position sensitivity during eye fixations was (means +/- S.D.) 1.8 +/- 0.9 spikes s-1 deg-1 for excitatory neurones and 2.2 +/- 1.3 spikes s-1 deg-1 for inhibitory neurones. Firing rate exhibited a high variability during eye fixations. Their responses during saccades in the off-direction were characterized by a pause that, although less defined, was occasionally present during saccades in the on-direction. Eye velocity sensitivity during spontaneous saccades in the on-direction was 0.17 +/- 0.15 spikes s-1 deg-1 s-1 for excitatory neurones and 0.15 +/- 0.07 spikes s-1 deg-1 s-1 for inhibitory vestibular neurones. During sinusoidal head stimulation at 0.2 Hz, vestibular neurones showed a type I discharge rate with a phase lead over eye position of 86.0 +/- 14.1 deg for excitatory and 80.2 +/- 12.5 deg for inhibitory neurones. Position sensitivity during vestibular stimulation did not differ significantly from values obtained for spontaneous eye movements. However, the velocity sensitivity of premotor vestibular neurones during head rotation was significantly higher (1.6 +/- 0.2 spikes s-1 deg-1 s-1 for excitatory and 1.5 +/- 0.3 spikes s-1 deg-1 s-1 for inhibitory neurones) than during spontaneous eye movements. 4. PH neurones projecting to the ABD nucleus were located in the rostral one-third of the nucleus. These neurones were excitatory on the ipsilateral and inhibitory on the contralateral ABD nucleus. Their firing rates were correlated mainly with eye position, increasing for abducting eye positions of the ipsilateral eye and decreasing with adduction movements.(ABSTRACT TRUNCATED AT 400 WORDS)

Conduction velocity of low-threshold mechanoreceptive afferent fibers in the glabrous and hairy skin of human hands measured with microneurography and spike-triggered averaging

The axonal conduction velocity (CV) of afferent fibers innervating low-threshold mechanoreceptors in the skin of the human hand was measured utilizing a spike-triggered averaging technique. Two tungsten microelectrodes were inserted into the median, or the ulnar, the superficial branch of the radial nerve, at two different points in the distal forearm. Unitary spike potentials were picked up with the proximal electrode. Unit-type SA (I and II), and FA (I and II), was determined from receptive field properties and response patterns to mechanical stimuli. Using these potentials as a trigger, the records from the distal electrode were averaged to reveal corresponding unitary potentials embedded in the background noise activities. CVs were calculated by dividing the interelectrode distance by the conduction time measured from the two neurograms. 122 mechanoreceptive afferent fibers in the glabrous and the hairy skin were recorded. The CVs of all sampled units were 36–73 m/s and the mean (SD) was 58.7 (7.4) m/s. The CVs did not differ between units in the three nerves, nor between units from the glabrous and the hairy skin. The mean CV of the FAI group was slower than the mean CV of the SA group by 4–5 m/s, but the overlap of the distributions was large.

Influence of synaptic identity on single-Ia-afferent connectivity and EPSP amplitude in the adult cat: homonymous versus heteronymous connections.

1. This study makes use of the pattern of synaptic connections between motoneurons and Ia afferents of triceps surae muscles in the cat to test the relative importance of synaptic identity, neuronal size, and neuronal topography as determinants of Ia-afferent connectivity and excitatory postsynaptic potential (EPSP) amplitude. 2. The synaptic actions of single-Ia medial gastrocnemius (MG) afferents were measured by intracellular recording in MG and lateral gastrocnemius (LG) motoneurons. The spike-triggered averaging technique was used to measure EPSPs generated by homonymous or heteronymous Ia afferents and motoneurons, i.e., neurons supplying the same or different muscles, respectively. In agreement with earlier studies, the pooled sample showed that the number of functional connections and the size of EPSPs were both significantly greater for homonymous than for heteronymous neurons. 3. Afferent conduction velocity, motoneuron conduction velocity, rheobase current, and position of the motoneuron relative to the spinal cord afferent entry were all correlated with EPSP amplitude, but the amplitude difference between homonymous and heteronymous connections remained significant after the statistical removal analysis of covariance (ANCOVA) of the contribution of these variables. Stepwise multiple-regression analysis showed that synaptic identity explained the greatest fraction of the variance in EPSP amplitude (9%), with significant but smaller fractions accounted for by rheobase current or motoneuron conduction velocity. 4. In a separate experiment, the monosynaptic affects from both homonymous and heteronymous single-Ia afferents were examined in each of 88 MG or LG motoneurons. The single-Ia afferents used in this portion of the study were sampled from both MG and LG muscles and selected for similar conduction velocities and spinal cord entry points.(ABSTRACT TRUNCATED AT 250 WORDS)

Evidence that the lateral tegmental field plays an important role in the central sympathoinhibitory action of 8-OH-DPAT

We examined the effects of 8-OH-DPAT and 5-HT on three types of sympathetic-related neurons identified in the lateral tegmental field of anesthetized cats using spike-triggered averaging techniques. Based on their response to baroreceptor activation, these neurons could be classified as sympathoexcitatory, sympathoinhibitory, or baroreceptor activation unresponsive. 8-OH-DPAT administered intravenously was neurons in a dose-dependent manner. Iontophoretic application of 8-OH-DPAT and 5-HT to the majority of sympathoexcitatory found to inhibit sympathoexcitatory neurons with a high degree of correlation to inhibition of sympathetic activity, and to excite sympathoinhibitory neurons caused inhibition of spontaneous activity while iontophoretic application of 8-OH-DPAT to sympathoinhibitory neurons had variable effects although 5-HT consistently caused excitation. Baroreceptor unresponsive neurons were insensitive to iontophoretic 8-OH-DPAT and showed only limited response to 5-HT. It is concluded that the lateral tegmental field plays an important role in the sympathoinhibitory action of 8-OH-DPAT.

Mono- and disynaptic pathways from Forel's field H to dorsal neck motoneurones in the cat.

1. We analysed the synaptic actions produced by Forel's field H (FFH) neurones on dorsal neck motoneurones and the pathways mediating the effects. 2. Stimulation of ipsilateral FFH induced negative field potentials of several hundred microvolts with the latency of about 1.1 ms in the medial ponto-medullary reticular formation, being largest in the ventral part of the nucleus reticularis pontis caudalis (NRPC), and in the dorsal part of the nucleus reticularis gigantocellularis (NRG). 3. Stimulation of ipsilateral FFH induced excitatory postsynaptic potentials (EPSPs) in 90% (47/52) and inhibitory postsynaptic potentials (IPSPs) in 19% (10/52) of the reticulospinal neurones (RSNs) in the NRPC and the NRG. Latencies of the EPSPs and IPSPs were 0.7-3.0 ms, the majority of which were in the monosynaptic range. The monosynaptic connexions were confirmed by spike triggered averaging technique both in excitatory (n = 4) and inhibitory (n = 2) pathways. 4. Single stimulation of FFH induced EPSPs at the segmental latencies of 0.3-1.0 ms in neck motoneurones, which were clearly in the monosynaptic range. Repetitive stimulation of FFH produced marked temporal facilitation of EPSPs in neck motoneurones. The facilitated components of the EPSPs had a little longer latencies and their amplitude reached several times as large as that evoked by single stimulation in all the tested motoneurones. These facilitated excitations are assumed to be mediated by RSNs in the NRPC and NRG, since RSNs were mono- and polysynaptically fired by stimulation of FFH and they were previously shown to directly project to neck motoneurones. 5. EPSPs were induced in 91% (82/91) of motoneurones supplying m. biventer cervicis and complexus (BCC; head elevator), 10% (3/29) of motoneurones supplying m. splenius (SPL; lateral head flexor). Likewise, stimulation of FFH produced EMG responses in BCC muscles, while not in SPL muscle. Thus FFH neurones produce excitations preferentially in BCC motoneurones. 6. Systematic tracking in and around FFH revealed that the effective sites for evoking above effects were in FFH and extended caudally along their efferent axonal course. 7. These results suggested that FFH neurones connect with neck motoneurones (chiefly BCC, head elevator) mono-, di- and/or polysynaptically and are mainly concerned with the control of vertical head movements.

Interaction of baseline synaptic noise and Ia EPSPs: evidence for appreciable negative correlation under physiological conditions

1. In the anesthetized cat, simultaneous intracellular recordings from pairs of spinal motoneurons were undertaken to see whether the amplitude of single-fiber excitatory postsynaptic potentials (EPSPs) in both cells fluctuated in a coordinated manner that would indicate correlative mechanisms at either pre- or post-synaptic level. Although these recordings revealed correlated fluctuations in the baseline, the single-fiber Ia/EPSPs recorded with the spike-triggered averaging technique exhibited no correlated fluctuations and, unexpectedly, virtually no increase in baseline variance associated with the EPSP. However, the fact that these experiments were carried out under conditions of high baseline synaptic noise (i.e., with muscle stretch) may have influenced the outcome because of interaction between EPSP and synaptic noise, and this possibility was evaluated explicitly. 2. A given connection was studied under low noise by electrically stimulating a single Ia fiber in the absence of muscle stretch. The same connection was analyzed under conditions of high noise by activating the fiber and all other stretch receptor afferents with muscle stretch and by using spike-triggered averaging to extract the EPSP. The differences in mean EPSP amplitude at a given connection under conditions of low noise and high noise were minimal. 3. Fluctuations in EPSP amplitude were then determined to see whether these were influenced by presence of baseline synaptic noise and whether the interaction was nonlinear. Two methods were used to measure EPSP fluctuations: measurement of the variance associated with the EPSP, and determination by the use of deconvolution methods of the discrete amplitude components associated with the EPSP. 4. An increase in baseline variance was observed during the EPSP evoked under low noise conditions at all six connections studied in this way. This increase disappeared at two of these connections when examined under high noise. This may help to explain the results obtained in pairs of motoneurons. 5. The deconvolution results were used to calculate the variance of the noise-free EPSP. This was found to differ from the variance of the EPSP amplitude distribution measured directly from the change in baseline variance associated with the EPSP. Analytic techniques suggested that this difference could be explained in most cases by negative correlation between the EPSP and baseline synaptic noise. These considerations led to an analytic method to assess the reliability of the deconvolution result. 6. Simulation studies revealed that the baseline variance increase associated with the EPSP is also highly dependent on the correlation between signal and noise.(ABSTRACT TRUNCATED AT 400 WORDS)

PREFERENTIAL GENERATION OF RECURRENT RESPONSES BY GROUPS OF MOTOR NEURONS IN MAN

Conventionally recorded surface F waves over the abductor digiti minimi muscle sampled a wide range of conduction velocities (CVs). Single motor unit (MU) F responses to threshold stimulation of the ulnar nerve were recorded with bipolar needle electrodes (BNE); the mean latencies of their fastest associated surface single unit M and F potentials were similar to those of the compound muscle action potential and fastest F wave, indicating that the fastest F wave indexes adequately the fastest motor fibre of the ulnar nerve. The mean surface unit F amplitude of MUs recorded by a BNE was similar to that of voluntary MUs recorded with a spike-triggered averaging technique; an estimated mean of 2-3 MUs per F wave was found by two methods. The frequency distribution of estimated F wave CVs was shifted towards faster values than expected from available studies of the distribution of CVs in single peripheral nerves; it was also higher than predicted from the expected relation between this distribution and the number of MUs per F wave, if an equal chance of activation and recording is assumed for each MU. There was a significant positive correlation between the frequency of F responses and their CV in 81 single MUs recorded by a BNE and tested with 200 threshold stimuli. These findings are consistent with preferential generation of recurrent responses by larger MUs; it may relate to a lesser chance of antidromic discharge in the smaller motor neurons and to a greater chance of collision of orthodromic (reflex) and antidromic impulses in their axons. The higher than expected percentage of all F waves that were repeater shapes or waves, and the presence of several distinct peaks in the distribution of intervals between repeaters of the same shape, suggest special, and heterogeneous, functional and anatomical arrangements in the groups of motor neurons generating them. Repeaters had greater amplitude and area, but similar latency and duration, than F wave shapes that did not repeat, suggesting that the former have a larger number of component MUs.

Relationship between electrical and mechanical properties of motor units.

In six healthy volunteers, the size of the motor unit action potential (MUAP) was related to its contractile behaviour. Electrical activity was determined by measuring amplitude and area of the potential, using the macro electromyography (EMG) technique. The mechanical response was characterised by contraction time, twitch force and relaxation time, recorded with the spike triggered averaging technique. With increasing tension of the whole muscle corresponding to an increasing threshold, the force of single units enlarged exponentially compared to their EMG. There was only a weak relationship between contraction time and electrical activity and none between EMG and relaxation time.

Vertical eye movement-related secondary vestibular neurons ascending in medial longitudinal fasciculus in cat. II. Direct connections with extraocular motoneurons

1. The preceding study in the alert cat has shown that many secondary vestibular axons that ascend in the medial longitudinal fasciculus (MLF) increase their firing rate in proportion to downward eye position. In the present study, projection and termination of these downward-position-vestibular (DPV) neurons within extraocular motoneuron pools were studied electrophysiologically by spike-triggered averaging techniques and morphologically be reconstructing their axonal trajectory after intra-axonal injection of horseradish peroxidase (HRP). 2. Extracellular field potentials recorded within the trochlear nucleus and/or the inferior rectus subdivision of the oculomotor nucleus were averaged by the use of spike potentials of single DPV neurons as triggers. All the crossed-DPV axons tested induced negative unitary field potentials in the trochlear nucleus (n = 9) and in the inferior rectus subdivision of the oculomotor nucleus (n = 5), suggesting that they made monosynaptic excitatory connection with motoneurons in these nuclei. The four crossed-DPV axons tested in the two motoneuron pools induced unitary field potentials in both. The majority of crossed-DPV axons terminated in these nuclei were directly activated from the caudal MLF, indicating that they had descending collaterals projecting to the spinal cord as well. The uncrossed-DPV axons did not induce such unitary field potentials either in the trochlear nucleus (n = 4) or in the inferior rectus subdivision (n = 3). 3. All the uncrossed-DPV axons examined (n = 14) induced positive unitary field potentials in the superior rectus subdivision of the oculomotor nucleus, suggesting that they made monosynaptic inhibitory connections with motoneurons innervating the superior rectus muscle. These uncrossed-DPV axons displayed regular firing patterns and were not activated from the caudal MLF. None of the crossed-DPV axons tested (n = 4) induced unitary field potentials in the superior rectus subdivision. 4. Five crossed-DPV axons were injected with HRP. All these axons ascended in the MLF contralateral to their soma, gave off many collaterals to the trochlear nucleus, and projected more rostrally. For three well-stained axons, numerous terminal branches were also found in the rostroventral part of the contralateral oculomotor nucleus, the area corresponding to the inferior rectus subdivision. Some collaterals in the oculomotor nucleus recrossed the midline to terminate in the medial part of the ipsilateral oculomotor nucleus. Other terminations were observed in the interstitial nucleus of Cajal and in the periaqueductal gray adjacent to the oculomotor nucleus. The crossed axons injected included both regular and irregular types, and three of the four examined were activated from the caudal MLF.(ABSTRACT TRUNCATED AT 400 WORDS)

Fatigue of single motor units in human masseter

The spike-triggered averaging technique was used to determine the time course and extent of fatigue of single motor unit twitches in the human masseter. This is the first report of a fatigue test having been applied to masseter motor units in either animals or humans. The human masseter was found to be comprised predominantly of fast-twitch motor units with a broad spectrum of fatigability. Very few physiological type S units were found, despite histochemical evidence for a substantial population of type I fibers in the masseter. In addition, there was no significant correlation between fatigability and either twitch amplitude or contractile speed in the motor units studied. The latter observations are consistent with the unusual histological features of the masseter. Comparison with other human fatigue data suggests that the extent of fatigue in the present population of masseter motor units after approximately 3,000 activations is similar to that reported for populations of units in first dorsal interosseous and medial gastrocnemius.

Inhibition of inspiratory neurons of the nucleus retroambigualis by expiratory neurons of the Botzinger Complex in the cat

The connection between expiratory neurons of the Botzinger Complex and contralateral inspiratory neurons of the nucleus retroambigualis was examined using the technique of spike-triggered averaging of intracellular potentials. Out of a total of 34 expiratory neurons found in the Botzinger Complex, 25 (73%) could be antidromically activated from the inspiratory region of the contralateral nucleus retroambigualis. The spike activities of 15 of these antidromically activated expiratory neurons were used as triggers for the averaging of the intracellular potentials recorded from 39 inspiratory neurons in the region of the contralateral nucleus retroambigualis. Unitary, inhibitory, postsynaptic potentials were observed in 11 of the 39 (28%) averages, from 6 of the 15 (40%) trigger neurons. It was concluded that these experiments demonstrate a monosynaptic inhibitory connection from expiratory neurons in the Botzinger Complex to inspiratory neurons in the contralateral nucleus retroambigualis.

Direct inhibitory synaptic connections of pontine omnipause neurons with burst neurons in Forel's field H related to vertical saccades in the cat

This study investigated synaptic connections of omnipause neurons (OPNs) in the midline pontine tegmentum with medium-lead burst neurons (BNs) in Forel's field H (FFH), using the spike-triggered averaging technique in chronically prepared alert cats. OPNs were identified behaviorally by tonic firing during the intersaccadic period, whereas medium-lead BNs were identified by spike bursts during upward or downward saccades. A positive shift of field potential in the BN area, associated with suppression of the BNs' bursts, was observed during tonic firing of OPNs and during microstimulation in the OPN area. Averaging of BN area field potentials triggered by spikes of single OPNs on both sides, which were antidromically evoked from the BN area, revealed positive waves with monosynaptic latencies. These results strongly suggest that OPNs on both sides make direct inhibitory synaptic connections with vertical eye movement-related medium-lead BNs in the FFH.

Effect of motor unit firing pattern on twitches obtained by spike‐triggered averaging

The spike-triggered averaging technique (STA) was used to determine the twitch profile of single motor units in human masseter during a voluntary isometric contraction. The effect of the immediate firing pattern of the unit on the twitch was assessed with a computer program which scanned the unit discharge records and selected valid trigger spikes for the averager on the basis of the interspike intervals preceding and following the trigger spike. Successive averages from the same data using different interval parameters revealed progressively more fusion of twitches as the instantaneous firing rate increased. When the data were averaged with interval parameters similar to those used in earlier studies, some fusion of the twitch profile was also evident. It is therefore likely that the degree of fusion of human motor unit twitches obtained by STA has been underestimated in the past. It is further concluded that masseter motor units are sufficiently fast-contracting to allow a relatively unfused twitch profile to be obtained with STA, provided trigger spikes are subjected to appropriate rate control.

Physiological properties of motor units in the human anterior digastric muscle

The activity of the single motor units in the human anterior digastric muscle was recorded in 7 subjects with a monopolar fine wire electrode. The force generated by jaw opening was simultaneously recorded with a mechanical transducer. The results obtained were as follows: (1) Increasing the force gradually, a few motor units were recruited orderly and the firing rate of each unit reached the peak frequency rapidly. These findings suggest that the control of the muscle force caused by jaw opening movements might depend more on the orderly recruitment of the motor units than on the rate modulation. (2) The contraction time, measured by the spike-triggered averaging technique varied from 25.6 to 132.4 ms. It was suggested that the anterior digastric muscle consisted of both the fast twitch motor units and the slow twitch motor units. (3) The motor units that have a higher threshold force tended to have a larger twitch tension and have a shorter twitch contraction time than that of the lower threshold motor units. This means that the anterior digastric muscle has a relatively simple role during the active jaw opening movements. (4) Fatigue can be estimated as the change in the firing rate. The motor units of a higher threshold force tended to be fatigue during the continuous jaw opening movements.

Acute effects of spinal transection on EPSPs produced by single homonymous Ia-fibers in soleus alpha-motoneurons in the cat.

1. Excitatory postsynaptic potentials (EPSPs) generated in soleus motoneurons by single homonymous Ia-fibers were measured using intracellular recording and the spike-triggered averaging technique. Two groups of barbiturate-anesthetized adult cats were studied: one with the spinal cord intact and the other with the spinal cord severed at thoracic segment 13 (T13) several hours prior to recording. 2. In cord-transected cats, single homonymous Ia-fibers produced EPSPs in soleus motoneurons that were, on average, larger and faster rising relative to normal, as they are for those produced in medial gastrocnemius (MG) motoneurons (8, 12, 13, 40). Specifically, mean EPSP amplitude and rise time were, respectively, 261 +/- 22 microV and 0.65 +/- 0.05 ms for the transected group vs. 160 +/- 21 microV and 0.96 +/- 0.08 ms for the intact group. The group means for each parameter were significantly different (P less than 0.005). 3. The group difference in EPSP amplitude was largely due to a decrease in number of small EPSPs in the transected group (11% less than 100 microV compared with the normal 41%) and not due to the occurrence of unusually large ones. Ratios of the largest to smallest amplitude EPSPs produced in the same motoneuron were similarly distributed for intact and transected groups, implying that the effect of transection on EPSP size was uniform across different Ia-fiber synapses made with the same motoneuron. Mean EPSP amplitude for each transected cat (n = 5) was larger than normal, but in some cases the increase took greater than 10 h to express itself. 4. The normal tendency for EPSP rise time to decline on average with amplitude was absent in the transected group, wherein rise time was reduced to similar average values in all amplitude categories. This suggests that the decrease in rise time occurred independently of the increase in amplitude. In contrast, EPSP half-width, which tended tow ward lower than normal values [5.63 +/- 0.36 (SE) ms vs. 6.51 +/- 0.44 ms; P greater than 0.10], decreased in proportion with rise time as evidenced by the preservation of the normal relation between those parameters in transected cats. Normalizing EPSPs by motoneuron time constant (tau) reduced the group differences in rise time and half-width, suggesting that a fall in tau contributes to the abbreviation of EPSP time course. 5. The condition of the spinal cord best accounted for differences in synaptic strength between groups.(ABSTRACT TRUNCATED AT 400 WORDS)

Monosynaptic projections of single muscle spindle afferents to neck motoneurons in the cat

Monosynaptic connections to dorsal neck motoneurons of the cat from single afferents supplying primary endings of neck muscle spindles were studied using spike-triggered averaging techniques. Single-fiber EPSPs were detected in only 11 of the 112 afferent-motoneuron pairs examined. The average amplitude of single-fiber EPSPs recorded in motoneurons with a membrane potential of greater than -40 mV was 49 microV. Motoneurons receiving functional contacts from a single afferent were confined to a small rostrocaudal zone within the motor nucleus. The low frequency of single-fiber EPSPs in neck motoneurons could not be attributed to the absence of projections to the ventral horn or to damage to either the afferents or motoneurons. Our results suggest, therefore, that single afferents from neck muscle spindles make functional contacts with a small fraction of neck motoneurons, unlike the arrangement seen in more commonly studied hindlimb muscle systems (Henneman and Mendell, 1981).

Projections from upper cervical inspiratory neurons to thoracic and lumbar expiratory motor nuclei in the cat

Previous studies have demonstrated the existence of axonal projections from propriospinal respiratory neurons in the rostral cervical cord of the cat (upper cervical inspiratory neurons) to the vicinity of the phrenic and rostral thoracic inspiratory (external) intercostal motoneurons. However, no synaptic targets of the upper cervical inspiratory neurons have been identified. This study investigated the axonal projections to the caudal thoracic and upper lumbar cord and the possible existence of inhibitory connections to the expiratory intercostal and abdominal motoneurons. The connections from upper cervical inspiratory neurons to expiratory motoneurons in the lower thoracic cord were examined using the methods of antidromic mapping and the spike-triggered averaging of intracellular potentials. Of the 70 upper cervical inspiratory neurons examined, only four (5.7%) could not be antidromically activated from the T9 segment of the spinal cord. The axons of 66 upper cervical inspiratory neurons were localized in the T9 or T10 segments of the thoracic spinal cord by antidromic activation at less than 5 μV, and the presence of collaterals was confirmed by antidromic mapping in 30 cases (49.2%). Of 21 axons tested for lumbar projections, 13 (61.9%) projected as far as T12, seven (33.3%) as far as L1, three (14.3%) as far as L2, and one (4.8%) was antidromically activated from L3. Spike-triggered averaging of the synaptic potentials recorded intracellularly from expiratory intercostal motoneurons in T9 and T10 spinal segments was done for 27 upper cervical inspiratory neurons, 17 of these with 4 or more motoneurons, for a total of 111 expiratory motoneurons. In 16 cases the motoneurons were injected with Cl − to reverse IPSPs and the spike-triggered averaging was repeated. No monosynaptic or disynaptic postsynaptic potentials were seen in any of the spike-triggered averages. We concluded that the upper cervical inspiratory neurons may provide inspiratory inhibition to expiratory motoneurons via a di- or oligosynaptic pathway involving segmental inhibitory interneurons and that either the spike-triggered averaging technique was not sensitive enough to detect the ipsilateral connections or the interneuron pathway was to the contralateral side of the spinal cord.