Serotonergic modulation of inspiratory hypoglossal motoneurons in decerebrate dogs

Inspiratory hypoglossal motoneurons (IHMNs) maintain upper airway patency. However, this may be compromised during sleep and by sedatives, potent analgesics, and volatile anesthetics by either depression of excitatory or enhancement of inhibitory inputs. In vitro data suggest that serotonin (5-HT), through the 5-HT2A receptor subtype, plays a key role in controlling the excitability of IHMNs. We hypothesized that in vivo 5-HT modulates IHMNs activity through the 5-HT2A receptor subtype. To test this hypothesis, we used multibarrel micropipettes for extracellular single neuron recording and pressure picoejection of 5-HT or ketanserin, a selective 5-HT2A receptor subtype antagonist, onto single IHMNs in decerebrate, vagotomized, paralyzed, and mechanically ventilated dogs. Drug-induced changes in neuronal discharge frequency (F(n)) and neuronal discharge pattern were analyzed using cycle-triggered histograms. 5-HT increased the control peak F(n) to 256% and the time-averaged F(n) to 340%. 5-HT increased the gain of the discharge pattern by 61% and the offset by 34 Hz. Ketanserin reduced the control peak F(n) by 68%, the time-averaged F(n) by 80%, and the gain by 63%. These results confirm our hypothesis that in vivo 5-HT is a potent modulator of IHMN activity through the 5-HT2A receptor subtype. Application of exogenous 5-HT shows that this mechanism is not saturated during hypercapnic hyperoxia. The two different mechanisms, gain modulation and offset change, indicate that 5-HT affects the excitability as well as the excitation of IHMNs in vivo.     

Intracellular analysis of respiratory-modulated hypoglossal motoneurons in the cat

Intracellular recordings from hypoglossal motoneurons in the brainstem of cats are described, along with postsynaptic potentials evoked by superior laryngeal, vagal and carotid sinus nerve stimulation. The study concentrates on hypoglossal motoneurons with respiratory-related discharge, which can be categorized into inspiratory, inspiratory/early-expiratory and expiratory patterns. Seven cells were labelled with horseradish peroxidase, their location and morphology are described. Stimulation of laryngeal receptors by balloon inflation or by water injection into the larynx, or mimicked by electrical stimulation of the superior laryngeal nerve results in enhanced postinspiratory activity in those cells (inspiratory/early-expiratory, expiratory) already receiving postinspiratory excitation; or actually produces a wave of postinspiratory depolarization in cells (inspiratory) previously quiescent during that period. It is concluded that the firing pattern of the respiratory-modulated hypoglossal motoneurons is unlikely to be static but depends on other factors, one of these being the level of ongoing, or previous laryngeal receptor stimulation.     

Connections between utricular nerve and dorsal neck motoneurons of the decerebrate cat.

1. We studied connections between the utricular (UT) nerve and dorsal neck motoneurons in decerebrate cats. Electrodes were fixed in place on the UT nerve under visual observation; the other branches of the vestibular nerve were transected. 2. The N1 field potential evoked by UT nerve stimulation was recorded in the vestibular nuclei at the start of each experiment. The potential typically grew until it reached a plateau. Stimulus spread (if any) to the central ends of other nerve branches was revealed by an additional increase in N1 amplitude after the plateau was reached. 3. We recorded intracellularly from 55 motoneurons in C1-C3. Some were identified as having axons in the dorsal rami, which innervate dorsal neck muscles. Others projected in nerves that were not available for stimulation. 4. UT nerve stimulation evoked synaptic potentials in essentially all motoneurons studied. The predominant pattern consisted of disynaptic excitatory postsynaptic potentials in ipsilateral motoneurons and inhibitory postsynaptic potentials that were at least trisynaptic in contralateral motoneurons. 5. The results demonstrate the presence of short-latency connections between the utricular nerve and dorsal neck motoneurons. The functional role of this pathway remains to be investigated.     

Mechanisms underlying excitatory effects of thyrotropin-releasing hormone on rat hypoglossal motoneurons in vitro.

1. The hypoglossal motor nucleus contains binding sites for the neuropeptide thyrotropin-releasing hormone (TRH) and is innervated by TRH-containing fibers. Although excitatory effects of TRH on hypoglossal motoneurons (HMs) have been described, the ionic mechanisms by which TRH exerts such effects have not been fully elucidated. Therefore, we investigated the effects of TRH on HMs in transverse slices of rat brainstem with intracellular recording techniques. 2. TRH was applied by perfusion (0.1-10 microM) or by pressure ejection (1.0 microM), while HMs were recorded in current or voltage clamp. In all cells tested, TRH caused a depolarization and/or the development of an inward current. These effects were fully reversible, dose dependent, and showed only modest desensitization with long applications. In addition, although TRH increased synaptic activity in many cells, the depolarizing response to TRH was maintained in tetrodotoxin (0.5-1.0 microM)-containing or in a nominally Ca(2+)-free perfusate containing 2 mM Mn2+. Thus TRH acts directly on HMs to cause the depolarization. 3. Hyperpolarizing current (or voltage) steps superimposed on the TRH-induced depolarization (or inward current) revealed a decreased input conductance. Extrapolated instantaneous current-voltage relationships obtained before and at the peak of the response to TRH intersected (i.e., reversed) at -101 mV, negative to the expected K+ equilibrium potential (EK). When extracellular [K+] was raised from 3 to 12 mM, the reversal potential was shifted in the depolarizing direction and the magnitude of the TRH-induced depolarization was diminished. Moreover, the TRH response was enhanced in size from depolarized potentials (i.e., further from EK). Taken together, these results indicate that TRH depolarizes HMs, in part, by decreasing a resting K+ conductance. 4. Similar to TRH, bath-application of 2 mM Ba2+ caused a depolarization associated with decreased conductance, suggesting that Ba2+ also blocks a resting K+ conductance. The Ba(2+)-sensitive and TRH-sensitive resting K+ conductances are apparently identical; in the presence of Ba2+, the customary TRH-induced decrease in conductance was occluded. 5. It is noteworthy that the TRH-induced inward current (ITRH), although diminished, was not entirely blocked by Ba2+. This second Ba(2+)-insensitive component of ITRH was not associated with a measurable change in input conductance. It was especially evident during current-clamp recordings, when the diminutive TRH-induced current was still capable of causing a substantial depolarization. The ionic basis of the residual TRH-induced inward current remains to be determined. 6. We investigated the functional consequences of these mechanisms of action of TRH on spike firing behavior of HMs.(ABSTRACT TRUNCATED AT 400 WORDS)     

Topographical arrangement of hypoglossal motoneurons: an HRP study in the cat

Myotopical localization of hypoglossal motoneurons and representation of the main branches of the hypoglossal nerve within the hypoglossal nucleus were examined in the cat by the HRP method. The hypoglossal nucleus is divided cytoarchitectonically into the ventromedial and dorsolateral divisions; the medial and lateral branches of the hypoglossal nerve are represented respectively in the ventromedial and dorsolateral divisions. The genioglossus motoneurons are located in the ventrolateral part of the ventromedial division, and the geniohyoid motoneurons are in the most ventral part of the ventromedial division. The hypoglossus and styloglossus motoneurons are located in the lateral and dorsolateral parts of the dorsolateral division.     

Excitatory and inhibitory postsynaptic potentials in cat hypoglossal motoneurons during swallowing

Summary The postsynaptic potentials produced in cat genioglossus and styloglossus motoneurons (GG- and SG-Mns) during swallowing were studied. During swallowing elicited by placing water on the dorsum of the tongue, the GG-muscle discharged for 80–210 ms (mean±S. D. 123±31 ms, N=59) and was abruptly suppressed, and the SG-muscle began discharging in synchrony with the GG-muscle and discharged for 200–360 ms (mean+ S. D. 247±36 ms, N=59). The GG and the SG-Mns were identified if unitary muscle activity followed the induced spike of the motoneuron one-for-one. During swallowing, excitatory postsynaptic potentials (EPSPs) were evoked in the SG-Mns regardless of the respiratory drive on the SG-Mns, and inhibitory postsynaptic potential (IPSP) or EPSP-IPSP was evoked on the GG-Mns regardless of the respiratory drive on the GG-Mns. By increasing the intracellular concentration of chloride ions, the IPSP elicited in the GG-Mn during swallowing was turned into a depolarizing potential. In immobilized cats, a depolarizing potential and a depolarizing-hyperpolarizing potential sequence was evoked successively on a tongue retractor motoneuron and a tongue protruder motoneuron by repetitive electrical stimulation of the superior laryngeal nerve.     

Corticofugal inhibitory effects on lingually induced postsynaptic potentials in cat hypoglossal motoneurons

The suppression of lingually or cortically induced postsynaptic potentials produced by conditioning stimulation of the cerebral cortex or the lingual nerve was studied in cat hypoglossal motoneurons. We have demonstrated that lingually or cortically induced inhibitory postsynaptic potentials were effectively suppressed by a conditioning stimulus of the cerebral cortex or the lingual nerve. In hypoglossal motoneurons after blocking inhibitory postsynaptic potentials by the administration of strychnine, lingually induced excitatory postsynaptic potentials and spikes were effectively suppressed by cortical stimulation. Whereas, a conditioning stimulus of the lingual nerve suppressed only a long-latency excitatory postsynaptic potential evoked by a test stimulus of the cerebral cortex, while a short-latency excitatory postsynaptic potential was unaffected. Picrotoxin and bicuculline appeared to act by reducing the suppression of lingually induced excitatory postsynaptic potentials produced by cortical conditioning stimulation.     

Postsynaptic potentials in the hypoglossal motoneurons set up by hypoglossal nerve stimulation.

In nembutalized cats intracellular potentials were recorded from hypoglossal motoneurons innervating either protruder or retractor muscles of the tonge (protruder and retractor motoneurons: P-Mns and R-Mns). Responses to stimulation of the hypoglossal nerve were explored and found to consist of an antidromic spike followed by an afterhyperpolarization (AHP) and a postsynaptic potential (PSP). When hypoglossal nerve stimulation was made with an intensity three times as large as the threshold for the hypoglossal motor fibers, the PSPs became evident under blockage of soma-dendritic invasion of the antidromic spike. In most of P-Mns or R-Mns, the PSPs were IPSPs, independent of the side of peripheral stimulation. The latencies were about 12 msec. Even when the cell membrane was hyperpolarized by injecting a hyperpolarizing current of up to 16 nA, the reversal point of the IPSP was difficult to find. In a small fraction of hypoglossal motoneurons the PSPs to hypoglossal nerve stimulation were EPSPs with latencies of 10 to 12 msec.     

Spike-firing resonance in hypoglossal motoneurons

During an inspiration the output of hypoglossal (XII) motoneurons (HMs) in vitro is characterized by synchronous oscillatory firing in the 20- to 40-Hz range. To maintain synchronicity it is important that the cells fire with high reliability and precision. It is not known whether the intrinsic properties of HMs are tuned to maintain synchronicity when stimulated with time-varying inputs. We intracellularly recorded from HMs in an in vitro brain stem slice preparation from juvenile mice. Cells were held at or near spike threshold and were stimulated with steady or swept sine-wave current functions (10-s duration; 0- to 40-Hz range). Peristimulus time histograms were constructed from spike times based on threshold crossings. Synaptic transmission was suppressed by including blockers of GABAergic, glycinergic, and glutamatergic neurotransmission in the bath solution. Cells responded to sine-wave stimulation with bursts of action potentials at low (<3- to 5-Hz) sine-wave frequency, whereas they phase-locked 1:1 to the stimulus at intermediate frequencies (3-25 Hz). Beyond the 1:1 frequency range cells were able to phase-lock to subharmonics (1:2, 1:3, or 1:4) of the input frequency. The 1:1 phase-locking range increased with increasing stimulus amplitude and membrane depolarization. Reliability and spike-timing precision were highest when the cells phase-locked 1:1 to the stimulus. Our findings suggest that the coding of time-varying inspiratory synaptic inputs by individual HMs is most reliable and precise at frequencies that are generally lower than the frequency of the synchronous inspiratory oscillatory activity recorded from the XII nerve.     

Firing rate modulation of motoneurons activated by cutaneous and muscle receptor afferents in the decerebrate cat

The aim of this study was to investigate whether activation of spinal motoneurons by sensory afferents of the caudal cutaneous sural (CCS) nerve evokes an atypical motor control scheme. In this scheme, motor units that contract fast and forcefully are driven by CCS afferents to fire faster than motor units that contract more slowly and weakly. This is the opposite of the scheme described by the size principle. Earlier studies from this lab do not support the atypical scheme and instead demonstrate that both CCS and muscle stretch recruit motor units according to the size principle. The latter finding may indicate that CCS and muscle-stretch inputs have similar functional organizations or that comparison of recruitment sequence was simply unable to resolve a difference. In the present experiments, we examine this issue using rate modulation as a more sensitive index of motoneuron activation than recruitment. Quantification of the firing output generated by these two inputs in the same pairs of motoneurons enabled direct comparison of the functional arrangements of CCS versus muscle-stretch inputs across the pool of medial gastrocnemius (MG) motoneurons. No systematic difference was observed in the rate modulation produced by CCS versus muscle-stretch inputs for 35 pairs of MG motoneurons. For the subset of 24 motoneuron pairs exhibiting linear co-modulation of firing rate (r > 0.5) in response to both CCS and muscle inputs, the slopes of the regression lines were statistically indistinguishable between the two inputs. For individual motoneuron pairs, small differences in slope between inputs were not related to differences in conduction velocity (CV), recruitment order, or, for a small sample, differences in motor unit force. We conclude that an atypical motor control scheme involving selective activation of typically less excitable motoneurons, if it does occur during normal movement, is not an obligatory consequence of activation by sural nerve afferents. On average and for both muscle-stretch and skin-pinch inputs, the motoneuron with the faster CV in the pair tended to be driven to fire at slightly but significantly faster firing rates. Computer simulations based in part on frequency-current relations measured directly from motoneurons revealed that properties intrinsic to motoneurons are sufficient to account for the higher firing rates of the faster CV motoneuron in a pair.     

An intracellular study of perineal and hindlimb afferent inputs onto sphincter motoneurons in the decerebrate cat

Summary The external urethral sphincter (EUS) and external anal sphincter (EAS) are striated muscles that function to maintain urinary and fecal continence respectively. This study examines the short-latency synaptic input from a variety of cutaneous perineal and muscle/cutaneous hindlimb afferents to the motoneurons innervating these muscles. Intracellular recordings from anti dromically identified EUS and EAS motoneurons provided records of the postsynaptic potentials (PSPs) produced by electrical stimulation of peripheral afferents in decerebrate or chloralose anesthetized cats. Excitatory postsynaptic potentials (EPSPs) were produced in most EUS and EAS motoneurons by stimulation of ipsilateral and contralateral sensory pudendal (SPud) and superficial perineal (SPeri) cutaneous nerves. The shortest cen tral latencies in the study (1.5 ms) suggest that there are disynaptic excitatory, in addition to tri-and oligosynap tic, connections within these reflex pathways. EPSPs mixed with longer latency inhibitory potentials (E/I PSPs) were observed in both motoneuron populations but were found more frequently in EAS motoneurons. These E/I PSPs were evoked more often from contralat eral afferents than from ipsilateral afferents. Cutaneous nerves innervating the hindlimb had weaker if any synaptic effects on sphincter motoneurons. Stimulation of ipsilateral hindlimb muscle nerves rarely produced PSPs in EUS motoneurons and had weak synaptic actions on EAS motoneurons. In 2 of 22 animals (both decerebrate), large inhibitory potentials predominated over early small EPSPs suggesting that inhibitory pathways from these afferents to sphincter motoneurons can be released under certain circumstances. The relation between the segmental afferents to EUS and EAS motoneurons and the neural circuitry influencing them during micturition and defecation are discussed.     

Membrane potential changes of phrenic motoneurons during fictive vomiting, coughing, and swallowing in the decerebrate cat.

1. The patterns of membrane potential changes of phrenic motoneurons were compared during fictive vomiting, fictive coughing, and fictive swallowing in decerebrate, paralyzed cats. These fictive behaviors were identified by motor nerve discharge patterns similar to those recorded from the muscles of nonparalyzed animals. Phrenic motoneurons (n = 54) were identified by antidromic activation from the thoracic phrenic nerve. Intracellular recordings were obtained from 27 motoneurons during fictive vomiting, 40 during fictive coughing, and 27 during fictive swallowing. Sixteen motoneurons were recorded during both fictive coughing and fictive swallowing, eight during both fictive coughing and fictive vomiting, and two during both fictive vomiting and fictive swallowing. Seven motoneurons were studied during all three behaviors. 2. Fictive vomiting, typically evoked by electrical stimulation of abdominal vagal afferents, was characterized by a series of bursts of coactivation of phrenic and abdominal motor nerves, culminating in an expulsion phase in which abdominal discharge was prolonged both with respect to phrenic discharge and to abdominal discharge during the preceding retching phase. During fictive vomiting, phrenic motoneurons depolarized abruptly, and the amplitude of depolarization was significantly greater than during control inspirations. They then repolarized slowly throughout the phrenic burst, rapidly repolarizing at the end of each phrenic burst during retching and reaching a level similar to that observed during expiration. During the expulsion phase, the pattern was initially the same. However, after the cessation of phrenic discharge, the membrane potential repolarized slowly until the end of the abdominal burst, exhibiting greater synaptic noise than during expiration. One phrenic motoneuron, presumably innervating the periesophageal region of the diaphragm, received a strong hyperpolarization just before the onset of the emetic episode and fired for shorter periods during fictive vomiting than did other phrenic motoneurons.(ABSTRACT TRUNCATED AT 250 WORDS)     

Responses of Renshaw cells coupled with hindlimb extensor motoneurons to sinusoidal stimulation of labyrinth receptors in the decerebrate cat

Contraction of ipsilateral limb extensors during side-down roll tilt of the head, leading to selective stimulation of labyrinth receptors, is attributed to an increased discharge of excitatory vestibulospinal (VS) neurons (alpha-responses) and a decreased discharge of medullary inhibitory reticulospinal (RS) neurons (beta-responses), both of which act on ipsilateral extensor motoneurons. Experiments were performed in decerebrate cats, with the de-efferented gastrocnemius-soleus (GS) muscle fixed at a constant length, to find out whether Renshaw (R) cells linked with GS motoneurons responded to labyrinth stimulation elicited by head rotation, while the neck had been bilaterally deafferented. We hoped in this way to clarify the role and the mechanism by which these inhibitory interneurons act on limb extensor motoneurons during the vestibular reflexes. 72.7% of the R-cells, disynaptically excited by group I volleys elicited by single shock stimulation of the GS nerve, weakly responded to head rotation at frequencies of 0.026-0.15 Hz and at a peak amplitude of 10 degrees. For the frequency of head rotation of 0.026 Hz, +/- 10 degrees C, most of the GS R-cells increased their firing rate during side-down head displacement (alpha-responses); some responses were related to head position, but others showed some phase lead or lag with respect to head position. The gain of the first harmonic of these unit responses was very low and corresponded on the average to 0.084 +/- 0.062, S.D. imp./s/deg, while the sensitivity corresponded to 2.14 +/- 2.35, S.D.%/deg (base frequency, 6.85 +/- 5.97, S.D. imp./s). These responses were attributed to the activity of VS neurons, the increased discharge of which during side-down head rotation exerts a weak excitatory influence on a limited number of GS motoneurons and, through their recurrent collaterals, on the related R-cells. The modulation of the firing rate of R-cells coupled with the GS motoneurons increased linearly by increasing the peak amplitude of displacement from 5 degrees to 20 degrees at the frequency of 0.026 Hz, so that the response gain remained almost unchanged. An increase in frequency of head rotation from 0.026 to 0.32 Hz at a fixed amplitude of 10 degrees, thus changing the maximal angular acceleration from 0.26 degrees/s2 to 41.7 degrees/s2, reversed the response pattern of R-cells reported above. The resulting beta-responses, which also showed some phase lead or lag with respect to head position, were attributed to vestibular activation of RS neurons.(ABSTRACT TRUNCATED AT 400 WORDS)     

Electrical properties of phrenic motoneurons in the cat: correlation with inspiratory drive

1. Resting membrane potential (Vmp), input resistance (Rn), rheobase (Irh), and after hyperpolarization duration (AHPdur) and amplitude (AHPamp) were measured in 38 phrenic motoneurons of anesthetized, paralyzed, and artificially ventilated cats during hypocapnic apnea. The mean +/- SD and range of values for these variables were as follows: Vmp, -68 +/- 5mV (range: -60 to -82); Rn, 1.3 +/- 0.6 M omega (0.6-2.4); Irh, 9.7 +/- 5 nA (2-20); AHPdur, 68 +/- 19 ms (37-134); AHPamp, 3.3 +/- 1.8 mV (1.0-8.5). In 31 motoneurons, the membrane potential level at which firing occurred (Vthr) during intracellular current injection was measured. The mean value of Vthr was -58 +/- 3 mV (range: -52 to -64). 2. A histogram of Rn revealed a bimodal distribution. Also a plot of Irh against Rn showed a grouping of the motoneurons into two subpopulations: 1) low-Rn and high-Irh cells, called type L neurons, and 2) high-Rn, low-Irh cells, called type H neurons. The overall negative linear correlation between Irh and Rn (r = -0.85; P less than 0.0001) resulted from this grouping rather than from a strictly linear relation between these two variables. 3. Electrical properties were compared for type L (n = 20) and type H (n = 18) phrenic motoneurons. The following mean values were found for each group, respectively: Rn, 0.8 and 1.8 M omega; Irh, 13.7 and 5.3 nA; AHPdur, 58 and 79 ms; AHPamp, 2.4 and 4.4 mV. All differences were significant (t test, P less than 0.001). Mean Vthr was the same for the two groups. 4. Comparison of these data with those available for lumbosacral motoneurons revealed that almost all investigated electrical properties of type L and type H phrenic motoneurons are similar to the analogous properties of type F (fast twitch) and type S (slow twitch) lumbosacral motoneurons, respectively. The apparent exception is the lower mean value of Irh for type L phrenic motoneurons compared with type F lumbosacral motoneurons. 5. For 13 cells, membrane potential was continuously monitored while spontaneous respiratory activity was restored by increasing CO2. It was found that at approximately the same end-tidal CO2 (about 7%) and a similar end-expiratory mean membrane potential level (approximately -70 mV), mean amplitude of peak inspiratory synaptic depolarization was higher in type H motoneurons (8.8 mV, n = 5) than in type L (2.9 mV, n = 8; P less than 0.001).(ABSTRACT TRUNCATED AT 400 WORDS)     

Variance analysis of excitatory postsynaptic potentials in cat spinal motoneurons during posttetanic potentiation

1. Fluctuations in the peak amplitudes of composite excitatory postsynaptic potentials (EPSPs) in cat spinal motoneurons were analyzed during posttetanic potentiation (PTP). Each of a series of identical tetanic stimulus trains delivered to a muscle nerve was followed by 45 test stimuli applied at 2-s intervals. The mean peak amplitude and mean peak variance were calculated for EPSPs evoked by all those stimuli following a tetanus with the same time interval. It was assumed that the variance arises primarily from the probabilistic all-or-none behavior of single synaptic boutons and background noise due to spontaneous synaptic activity and thermal noise in the recording system. The variance was corrected for the contribution from additive Gaussian background noise. 2. If it is assumed that individual synaptic boutons behave independently, corrected mean peak variance and mean peak amplitude are related by a parabolic function. The expected parabolic relationship was seen in 9 of 31 cases studied, and the parameters of the best parabolic fit to the data allowed estimation of some synaptic properties. From these parameters, the mean amplitude of the unit EPSP (v) was estimated to be 102.1 +/- 57.4 (SD) microV. An average of 3.7 boutons comprised each Ia-motoneuron contact system. 3. On average, only 27% of all synaptic boutons given off by the stimulated Ia fibers to one motoneuron were active and releasing transmitter during unpotentiated reflex transmission. The remaining 73% of the synapse population was intermittently silent. The population of boutons which took part in synaptic transmission could be divided into two subpopulations, one with a release probability P = 1 and a second with a mean release probability P = 0.13 +/- 0.086. 4. We conclude that synaptic boutons connecting Ia afferents to motoneurons exist in two populations, one having a high and one a low probability of transmitter release. Transmitter release is quantal, resulting in a unit EPSP of approximately 100 microV measured at the motoneuron soma.     

Synaptic efficacy of inhibitory synapses in the reinnervating hypoglossal motoneurons.

The synaptic efficacy of inhibitory synapses in tongue protruder motoneurons reinnervating the tongue retractor muscle was studied in cats. We have demonstrated that the percentage magnitude of a short- and a long-lasting inhibitory postsynaptic potential in the inhibitory postsynaptic potentials produced in the tongue protruder motoneurons, whose axons had been cut but allowed to regenerate to make functional contact with the tongue retractor muscles, by lingual nerve or inferior alveolar nerve stimulation, was rearranged to appear like that exhibited by the tongue retractor motoneurons that normally supply that muscle. In addition, the peak amplitude of the summated afterhyperpolarization in a tongue protruder motoneuron on operated cats at nine months axon-union was in the normal range.     

The distribution of inhibitory and excitatory synapses on single, reconstructed jaw-opening motoneurons in the cat

In a previous study, we reported that the distribution of inhibitory input, in contrast to excitatory input, decreased somatofugally along dendrites of cat jaw-closing alpha-motoneurons [J Comp Neurol 414 (1999) 454]. The present study examined the distribution of GABA, glycine, and glutamate immunopositive boutons covering horseradish peroxidase-labeled cat jaw-opening motoneurons. The motoneurons were divided into four compartments: the soma, and primary, intermediate, and distal dendrites. Ninety-seven percent of the total number of studied boutons had immunoreactivity for at least one of the three amino acids. The proportion of boutons immunoreactive for GABA and/or glycine was lower than the proportion of boutons immunoreactive for glutamate. Boutons immunoreactive to glycine alone were more numerous than boutons double-labeled for GABA and glycine, which, in turn, occurred more frequently than boutons immunoreactive to GABA alone. The percentage synaptic covering (proportion of membrane covered by synaptic boutons) of the putatively excitatory (glutamate containing) and putatively inhibitory (GABA and/or glycine containing) boutons decreased somatofugally along the dendrites. Such systematic variations were not seen in the packing density (number of boutons per 100 microm(2)); the packing density showed a distinct drop between the soma and primary dendrites but did not differ significantly among the three dendritic compartments. Overall, the packing density was slightly higher for the putatively excitatory boutons than for the inhibitory ones. When taken together with previous analyses of jaw-closing alpha-motoneurons the present data on jaw-opening alpha-motoneurons indicate that the two types of neuron differ in regard to the nature of synaptic integration in the dendritic tree.