Mossy and climbing fiber activity during phasic and tonic phenomena of sleep

Summary In the intact, unanesthetized, freely-moving cat the Purkinje cell activity driven by the mossy fiber system presents phasic increases in coincidence with movements of wakefulness or desynchronized sleep. Its rate of discharge may range between wide limits.In contrast, activity driven by the climbing fiber system varies between much more restricted limits. In the absence of movements its rate of dischange is higher during the desynchronized stage of sleep as compared to the synchronized one. In coincidence with the phasic events of paradoxical sleep the rate of discharge presents often a decrease. Some units, which have been assumed to be mossy and climbing fiber afferents, have a behavior similar to the mossy and climbing fiber driven activity, cespectively. Some characteristics of olivo-cerebellar neurons have been presented.A possible role of the mossy fiber and climbing fiber systems, respectively for phasic and tonic functions, is discussed. The pattern of activity during sleep is discussed in relation to the pattern observed in other brain areas.It is suggested that the cerebellum contributes to the collapse of the postural tonus which occurs during paradoxical sleep.     

A cholinoceptive desynchronized sleep induction zone in the anterodorsal pontine tegmentum: spontaneous and drug-induced neuronal activity

The effect of carbachol microapplication (4 micrograms/250 nl per 90 s) on the discharge of neurons in the anterodorsal pons of four cats was studied using a newly devised microinjector-microelectrode assembly. Neurons were classified according to the magnitude of their discharge rate increases (or decreases) in physiological desynchronized sleep as desynchronized-on (or desynchronized-off) before injecting carbachol. When carbachol produced a desynchronized sleep-like state only half (15 out of 30) of the desynchronized-on cells were activated (desynchronized-on/desynchronized sleep-like state-on) while the other half were not (desynchronized-on/desynchronized sleep-like state-not on). Compared with the non-activated cells, the desynchronized-on/desynchronized sleep-like state-on cells had three features consistent with playing an active role in desynchronized sleep generation: these cells had a higher mean discharge frequency in desynchronized sleep and higher ratio of discharge frequency in desynchronized sleep compared with wakefulness; they did not fire in phase with electromyogram excitation of neck muscles; and they were concentrated in the short latency desynchronized sleep-like state induction zone described in the companion paper. The three-way correlation between the optimal anatomical site for short latency desynchronized sleep-like state induction, the selective neuronal discharge pattern in desynchronized sleep and the cholinergic activation pattern in the desynchronized sleep-like state suggest that we may have identified a neuronal population that is cholinoceptively activated as part of the physiological mechanism of desynchronized sleep generation.     

Unitary characteristics of presumptive cholinergic tegmental neurons during the sleep-waking cycle in freely moving cats

Summary A total of 260 neurons were recorded in the rostral pontine tegmentum of freely moving cats during the sleep-waking cycle. Of these, 207 neurons (80%) were located in the dorsal pontine tegmentum containing monoaminergic and choline acetyltransferase (ChAT)-immunoreactive, or cholinergic neurons. In addition to presumably monoaminergic PS-off cells (n = 51) showing a cessation of discharge during paradoxical sleep (PS) and presumably cholinergic PGO-on cells (n = 40) exhibiting a burst of discharge just prior to and during ponto-geniculo-occipital (PGO) waves, we observed tonic (n = 108) and phasic (n = 61) neurons exhibiting, respectively, tonic and phasic patterns of discharge during wakefulness and/or paradoxical sleep. Of 87 tonic cells histologically localized in the dorsal pontine tegmentum rich in cholinergic neurons, 46 cells (53%) were identified as giving rise to ascending projections either to the intralaminar thalamic complex (n = 26) or to the ventrolateral posterior hypothalamus (n = 13) or to both (n = 9). Two types of tonic neurons were distinguished: 1) tonic type I neurons (n = 28), showing a tonic pattern and high rates of discharge during both waking and paradoxical sleep as compaired with slow wave sleep; and 2) tonic type II neurons (n = 20), exhibiting a tonic pattern of discharge highly specific to the periods of paradoxical sleep. Tonic type I neurons were further divided into two subclasses on the basis of discharge rates during waking: a) rapid (Type I-R; n = 17); and b) slow (Type I-S; n = 11) units with a discharge frequency of more than 12 spikes/s or less than 5 spikes/s, respectively. Like monoaminergic PS-off and cholinergic PGO-on cells, both tonic type II and type I-S cells were characterized by a long spike duration (median: 3.3 and 3.5 ms), as well as by a slow conduction velocity (median: 1.8 and 1.7 m/s). In the light of these data, we discuss the possible cholinergic nature and functional significance of these ascending tonic neurons in the generation of neocortical electroencephalographic desynchronization occurring during waking and paradoxical sleep.     

Discharge patterns of human genioglossus motor units during sleep onset

STUDY OBJECTIVES: Multiunit electromyogram recordings of genioglossus have demonstrated an abrupt reduction in the muscle's activity at sleep onset. Recent evidence from single motor unit recordings indicates that the human genioglossus muscle consists of motor units with a variety of discharge patterns. The aim of the present study was to determine the effect of sleep onset on the activity of individual motor units as a function of their particular discharge pattern. DESIGN: Genioglossus activity was assessed using intramuscular fine-wire electrodes via a percutaneous approach. Sleep onsets (alpha-to-theta transitions) were identified and the genioglossus electromyogram recordings analyzed for single motor unit activity. SETTING: Sleep research laboratory. PARTICIPANTS: Sleep and respiratory data were collected in 8 healthy subjects (6 men). MEASUREMENTS AND RESULTS: One hundred twenty-seven motor units were identified: 23% inspiratory phasic, 45% inspiratory tonic, 4% expiratory phasic, 9% expiratory tonic, 16% tonic, and 3% other. Approximately 50% of inspiratory units (phasic and tonic) ceased activity entirely at sleep onset, whereas those inspiratory units that continued to be active showed a reduction in the proportion of each breath over which they were active. However, the rate of discharge of inspiratory units during the period they did fire was not altered. In contrast, tonic and expiratory units were unaffected by sleep onset, maintaining their discharge pattern over the alpha-to-theta transition. CONCLUSIONS: Central control of inspiratory motoneuron output differs from that of tonic and expiratory units during sleep onset, suggesting that the maintenance of airway patency during sleep may become more reliant on the stiffening properties of tonic and expiratory modulated motor units.     

Paradoxical sleep and its chemical/structural substrates in the brain

As originally named for the ostensibly contradictory appearance of rapid eye movements and low voltage fast cortical activity during behavioral sleep, paradoxical sleep or rapid eye movement sleep, represents a distinct third state, in addition to waking and slow wave sleep, in mammals and birds. It is an internally generated state of intense tonic and phasic central activation that is contemporaneous with the inhibition of sensory input and motor output.

In early studies, it was established that the state of paradoxical sleep was generated within the brainstem, and particularly within the pons. Pharmacological studies indicated an important role for acetylcholine as a neurotransmitter in the generation of this state. Local injections of cholinergic agonists into the pontine tegmentum triggered a state of paradoxical sleep marked by phasic ponto-geniculo-occipital spikes in association with cortical activation and neck muscle atonia. Following the immunohistochemical identification of choline acetyl transferase-containing neurons and their localization to the dorsolateral ponto-mesencephalic tegmentum, neurotoxic lesions of this major cholinergic cell group could be performed to assess its importance in paradoxical sleep. Destruction of the majority of the cholinergic cells, which are concentrated within the laterodorsal tegmental and pedunculopontine tegmental nuclei but extend also into the locus coeruleus and parabrachial nuclei in the cat, resulted in a loss or diminishment of the state of paradoxical sleep, ponto-geniculo-occipital spiking and neck muscle atonia. These deficits were correlated with the loss of choline acetyltransferase-immunoreactive neurons in the region, so as to corroborate results of pharmacological studies and single unit recording studies indicating an active role of these cholinergic cells in the generation of paradoxical sleep and its components. These cells provide a cholinergic innervation to the entire brainstem reticular formation that may be critical in the generation of the state which involves recruitment of massive populations of reticular neurons. Major ascending projections into the thalamus, including the lateral geniculate, may provide the means by which phasic (including ponto-geniculo-occipital spikes) and tonic activation is communicated in part to the cerebral cortex. Descending projections through the caudal dorsolateral pontine tegmentum and into the medial medullary reticular formation may be involved in the initiation of sensorimotor inhibition.

Although it appears that the pontomesencephalic cholinergic neurons play an important, active role in the generation of paradoxical sleep, this role may be conditional upon the simultaneous inactivity of noradrenaline and serotonin neurons, evidence for which derives from both pharmacological and recording studies. An interaction between the cholinergic and noradrenergic (as well as serotoninergic) neurons may potentially occur within the dorsolateral pontine tegmentum where the soma and processes of the two cell types are in close proximity, but also by the potential intermediary of local GABA-containing neurons that would permit a mutually inhibitory relationship between the two through the sleep cycle.

The cyclic activation of reticular neurons during paradoxical sleep may release intrinsic brainstem patterns of sensorimotor processes and complex behaviors that underlie oneiric behavior and dreams. These fundamental processes may be important for the programming of the central nervous system during development, when paradoxical sleep is predominant, as well as for reinforcing and modulating these behaviors through life.     

Firing rates and patterns of midbrain reticular neurons during steady and transitional states of the sleep-waking cycle

Summary Spontaneous firing of midbrain reticular formation (MRF) neurons was recorded extracellularly in chronically implanted, behaving cats during steady and transitional states of the sleep-waking cycle. Physiological identification of receiver and/or projection MRF neurons was achieved through orthodromically and antidromically elicited discharges.Discharge rates of MRF neurons were more than double in waking (W) and active sleep (D) without phasic motor events, as compared to synchronized sleep (S). During behavioral states associated with EEG activation, the increased firing was essentially due to cells exhibiting high discharge rates, located at relatively ventral levels of the midbrain core. MRF neurons with identified rostrally projecting axons were more active during W and D states; their discharge rates were significantly higher than those of caudally projecting cells. The discharge patterns of MRF neurons increasing their firing rates from S to W and D were of the tonic type. First-order analyses showed a negligible proportion of both very short and long interspike intervals in all states, large interval density around the mode especially in W and D, and the smallest variation coefficients in W. Rhythmic firing with a period near the modal interval was detected during W by autocorrelations.The increase in firing rate of MRF neurons from S to W or D took place before overt EEG desynchronization and behavioral manifestations that define stable W or D states. In our sample a statistically significant increase in discharge rate was found about 15 s before the end of S sleep epochs that developed into awakening.The differences between discharge features of MRF neurons during waking and sleep states and those of neurons in other brainstem reticular fields are emphasized. Taken together, these data support, at a cellular level, Moruzzi and Magoun's concept of a rostral reticular substrate that gives rise to impulses leading to tonic activation of the thalamocortical systems.Supported by grant MT-3689 from the Medical Research Council of CanadaPart of the results was incorporated in the Ph. D. thesis of N. Ropert     

Discharge Patterns of Human Genioglossus Motor Units during Arousal from Sleep

Study Objectives:

Single motor unit recordings of the human genioglossus muscle reveal motor units with a variety of discharge patterns. Integrated multiunit electromyographic recordings of genioglossus have demonstrated an abrupt increase in the muscle''s activity at arousal from sleep. The aim of the present study was to determine the effect of arousal from sleep on the activity of individual motor units as a function of their particular discharge pattern.

Design:

Genioglossus activity was measured using intramuscular fine-wire electrodes inserted via a percutaneous approach. Arousals from sleep were identified using the ASDA criterion and the genioglossus electromyogram recordings analyzed for single motor unit activity.

Setting:

Sleep research laboratory.

Participants:

Sleep and respiratory data were collected in 8 healthy subjects (6 men).

Measurements and Results:

138 motor units were identified during prearousalarousal sleep: 25% inspiratory phasic, 33% inspiratory tonic, 4% expiratory phasic, 3% expiratory tonic, and 35% tonic. At arousal from sleep inspiratory phasic units significantly increased the proportion of a breath over which they were active, but did not appreciably increase their rate of firing. 80 new units were identified at arousals, 75% were inspiratory, many of which were active for only 1 or 2 breaths. 22% of units active before arousal, particularly expiratory and tonic units, stopped at the arousal.

Conclusions:

Increased genioglossus muscle activity at arousal from sleep is primarily due to recruitment of inspiratory phasic motor units. Further, activity within the genioglossus motoneuron pool is reorganized at arousal as, in addition to recruitment, ∼20% of units active before arousals stopped firing.

Citation:

Wilkinson V; Malhotra A; Nicholas CL; Worsnop C; Jordan AS; Butler JE; Saboisky JP; Gandevia SC; White DP; Trinder J. Discharge patterns of human genioglossus motor units during arousal from sleep. SLEEP 2010;33(3):379-387.     

Bulbo-thalamic neurons related to thalamocortical activation processes during paradoxical sleep

Summary Neurons histologically localized in the gigantocellular (Gc) and magnocellular (Mc) fields of the bulbar reticular formation were tested for antidromic invasion by stimulating the ventromedial (VM) and intralaminar (centralis lateralis, CL, and centrum medianum, CM) thalamic nuclei, midbrain reticular formation (MRF), and reticulospinal tract. An overwhelming majority (94%) of antidromically identified cells projected either to rostral structures (MRF, medial and intralaminar thalamic nuclei) or to the spinal cord, while only 6% had bifurcating axons.Rostrally projecting bulbar reticular neurons were investigated during various wake-sleep behavioral states, (a) Phasic neurons were related to PGO waves, eye and head movements, and were localized in both Gc and Mc fields, (b) The majority of tonic neurons projected to MRF and VM and they were localized within Mc in a proportion of 85%. In order to test their possible role in activation of thalamocortical processes (as betrayed by EEG desynchronization), the activity of tonically discharging cells was separately evaluated in periods with and without phasic motor events. Half of the tonically discharging neurons had a high selectivity of discharge during paradoxical sleep without REM bursts (PS-); the ratio of their mean discharge rate during PS- to that in quiet wakefulness (QW) or slow-wave sleep (SWS) was 8 and 6, respectively. The other half of the tonic neurons equally increased firing rates from SWS to either QW or PS.The firing rate of rostrally projecting bulbar reticular neurons with tonic discharge patterns was analyzed during transitions from SWS to PS. An increase in discharge rate was found about 30 to 60 s prior to the first sign of EEG desynchronization in PS, during fully synchronized sleep with PGO waves (S-PGO). Statistical testing showed that the increased firing rate was not associated to PGO waves, but was temporally related to the appearance of EEG desynchronization at PS onset. We conclude, on the basis of these and other recent data, that tonically discharging bulbar reticular neurons with identified projections to the midbrain and thalamic nuclei act synergically with rostrally projecting MRF neurons as sources of thalamocortical activation.Supported by grant MT-3689 from Medical Research Council of CanadaSupported by INSERM (U 52), CNRS (LA 162) and DRET (grant 81-205)     

大鼠低位脑干睡眠相关结构胆碱能神经元的分布

本文用胆碱乙酰转移酶的单克隆抗体免疫组织化学技术,观察了胆碱能神经元在大鼠低位脑干睡眠相关结构的分布。结果表明,蓝斑是非胆碱能的,很蓝斑腹侧部网状结构含有胆碱乙酰转移酶阳性反应神经元。胆碱乙酰转移酶还出现在脑桥内侧网状结构,相应于脑桥尾侧网状核以及中缝大核。延髓网状巨细胞核及其腹侧部也有胆碱能神经元出现。这些区域的胆碱能神经元可能参与异相睡眠的诱发及其某些特性的产生,也可能和慢波睡眠有关。     

Postnatal development of spontaneous neuronal discharges in the pontine reticular formation of free-moving rats during sleep and wakefulness

Summary A developmental study has been made of spontaneous neuronal activity within the pontine reticular formation (giant cell field: FTG) of the rat between one week and one month after birth. Through day 14, the recorded FTG neurons discharged more frequently during quiet sleep (QS) than was generally true in older animals. In addition, they were active to the same extent during active-sleep (AS) as during waking-with-movements (AW). In contrast, most of the cells recorded from day 15 on were considerably more active during AS and AW, relative to the QS level, than had hitherto been the case. This new class of neurons, in turn, fell into two sub-groups, one of which was most active during AW while the other was more active during AS. Clomipramine selectively suppressed AS along with the neuronal activity patterns associated with it, and in many cases the QS firing level was even lower than it had been prior to the injection. It is concluded that FTG unit activity is an excellent monitor for controlling the effectiveness of experimental manipulations of AS but is probably not involved in its generation.     

The regulation of heart rate during sleep

Summary One group of the experiments was carried out on cats using encéphale isolé preparations, a second group was performed on cats with chronically implanted electrodes. Cortical EEG, electromyogram, ocular movements and electrocardiogram were recorded simultaneously.Different kinds of heart denervation have been performed in order to study separately the action of the sympathetic and parasympathetic nervous system. The following results have been obtained:The changes in heart rate associated with the different states of sleep and wakefulness are controled by both the sympathetic and parasympathetic section of the autonomic nervous system.The tonic fall in heart rate during a shift from wakefulness to synchronized sleep is prepondantly caused by an increase in the vagal tonus, whereas the further decrease in heart rate during desynchronized sleep is mainly brought about by a tonically reduced sympathetic discharge.The phasic increase in heart rate following an arousal is only due to a phasic inhibition of the parasympathetic cardioinhibitory center. The phasic changes in heart rate associated with the bursts of rapid eye movements occuring during desynchronized sleep are caused by changes in the sympathetic and vagal activity as well. A detailed analysis shows that the phasic acceleration is prepondantly due to an inhibition of the vagal activity, whereas the following bradycardia is brought about by a phasic inhibition of the sympathetic output and a simultaneous increase in the vagal discharge.The results clearly demonstrate that the changes in heart rate related with the different states of sleep and wakefulness are not caused by quantitatively equal antagonistic changes in the activity of the two sections of the autonomic nervous system.This work was supported by grants from the Deutsche Forschungsgemeinschaft.     

Independent and convergent signals from the pontomedullary reticular formation contribute to the control of posture and movement during reaching in the cat

We have addressed the nature of the postural control signals contained within the discharge activity of neurons in the pontomedullary reticular formation, including reticulospinal neurons, during a reaching task in the cat. We recorded the activity of 142 neurons during ipsilateral reaching movements that required anticipatory postural adjustments (APAs) in the supporting limbs to maintain equilibrium. Discharge activity in 82/142 (58%) neurons was significantly increased before the onset of the reach. Most of these neurons discharged either in a phasic (22/82), tonic (10/82), or phasic/tonic (41/82) pattern. In each of these 3 groups, the onset of the discharge activity in some neurons was temporally related either to the go signal or to the onset of the movement. In many neurons, one component of the discharge sequence was better related to the go signal and another to the onset of the movement. Based on our previous behavioral study during the same task, we suggest that reticular neurons in which the discharge activity is better related to the go signal contribute to the initiation of the APAs that precede the movement. Neurons in which the discharge activity is better related to the movement signal might contribute to the initiation of the movement and to the production of the postural responses that accompany that movement. Together our results suggest the existence of neurons that signal posture and movement independently and others that encode a convergent signal that contributes to the control of both posture and movement.     

Enhancement of dorsal raphe discharge by medial pontine reticular formation stimulation depends on behavioral state

Single cell activity was recorded with microwire electrodes from the dorsal raphe nucleus (DRN) of cats across multiple cycles of sleep and wakefulness. Electrical stimulation of the medial pontine reticular formation (mPRF) produced a 10-fold enhancement of DRN discharge during wakefulness and slow wave sleep but not during desynchronized sleep. The neuronal mechanisms likely to mediate the state-dependent effects of mPRF stimulation are discussed with regard to the hypothesis that DRN plays a major role in behavioral state control.     

Elimination of paradoxical sleep by lesions of the pontine gigantocellular tegmental field in the cat.

Bilateral lesions of the pontine gigantocellular tegmental field in the cat resulted in the complete elimination of paradoxical sleep during 3 weeks postoperative recording. The tonic muscular atonia, normally characteristic of this state, was absent. The phasic components, rapid eye movements and ponto-geniculooccipital (PGO) spikes, did not occur in association with an activated EEG, as they normally do in paradoxical sleep. In fact, PGO spikes were virtually absent immediately after the lesion and were only secondarily apparent as isolated phenomena during slow wave sleep to represent in total daily number 5% of normal the first week and 15% of normal the third week after the lesion. These results indicate that neurons whose perikarya and/or processes are located within the pontine gigantocellular tegmental field and which are not part of the noradrenaline locus coeruleus complex, are critical for paradoxical sleep.     

Tonic inhibition and ponto-geniculo-occipital-related activities shape abducens motoneuron discharge during REM sleep

Eye movements, ponto-geniculo-occipital (PGO) waves, muscular atonia and desynchronized cortical activity are the main characteristics of rapid eye movement (REM) sleep. Although eye movements designate this phase, little is known about the activity of the oculomotor system during REM sleep. In this work, we recorded binocular eye movements by the scleral search-coil technique and the activity of identified abducens (ABD) motoneurons along the sleep–wake cycle in behaving cats. The activity of ABD motoneurons during REM sleep was characterized by a tonic decrease of their mean firing rate throughout this period, and short bursts and pauses coinciding with the occurrence of PGO waves. We demonstrate that the decrease in the mean firing discharge was due to an active inhibition of ABD motoneurons, and that the occurrence of primary and secondary PGO waves induced a pattern of simultaneous but opposed phasic activation and inhibition on each ABD nucleus. With regard to eye movements, during REM sleep ABD motoneurons failed to codify eye position as during alertness, but continued to codify eye velocity. The pattern of tonic inhibition and the phasic activations and inhibitions shown by ABD motoneurons coincide with those reported in other non-oculomotor motoneurons, indicating that the oculomotor system – contrary to what has been accepted until now – is not different from other motor systems during REM sleep, and that all motor systems are receiving similar command signals during this period.     

Impulse activity of ventral tegmental area neurons during heroin self-administration in rats

To assess the pattern of mesocorticolimbic dopamine activity associated with drug-seeking and drug-taking behavior, we recorded impulse activity of ventral tegmental area neurons during intravenous heroin self-administration in trained rats. Although these neurons had considerable variability, two major groups-units with triphasic long-duration spikes and biphasic short-duration spikes-were identified. Relative to a slow and irregular basal activity of long-spike units, the first self-administration of each session was preceded by a phasic neuronal activation and followed by a more sustained drug-induced activation that reached a maximum at the time of the second self-injection. After each subsequent heroin self-injection, the discharge rate transiently decreased, correlating with the blockade of preceding motor activation and the appearance of freezing, but slowly and gradually increased again in parallel with searching behavior, reaching a maximum at the time of the next self-injection. Passive drug injections in either drug-naive, freely moving or drug-experienced, anesthetized rats caused much smaller, tonic increases in activity of long-spike units; these monophasic increases changed into biphasic responses with repeated injections. Although short-spike units had highly varying discharge rate and showed phasic activation during movement, during heroin self-injections they generally mimicked the activity pattern seen in long-spike units. Our results indicate that in behaving animals indirect "identification" of dopamine cells based on their distinctive electrophysiological features is more complex than in vitro and in anesthetized preparations. With respect to long-spike units, a candidate group of presumed dopamine neurons, our data agree with the view that mesocorticolimbic dopamine activation is important for the activational and/or motivational aspects of heroin-taking behavior and suggest the role of an abrupt termination of dopamine activation for drug reinforcement (reward). Although the neurochemical nature of long- and short-spike units is obviously different, similar changes in their activity may indicate that they are regulated by similar afferent inputs and that these inputs change similarly during drug-taking behavior.     

Modulation of unit activity in the amygdala of unrestrained cats during the sleep-waking cycle

Recordings of amygdaloid unit activity were performed in unrestrained cats during the sleep-waking cycle. Discharge rates of most units studied in the central part of the amygdala were related to the sleep-wakefulness cycle. Out of 30 units, 18 had decreased discharge rates during slow-wave-sleep (SWS), compared to the waking state. In 12 of these 18 units discharge rates became higher during desynchronized sleep (DS). A similar relationship was seen when studying units in basal amygdala.     

Distribution of choline acetyltransferase immunoreactive somata in the feline brainstem: implications for REM sleep generation

In the present study we examined the distribution of cholinergic and catecholaminergic neurons, in the feline brainstem, as defined by choline acetyltransferase (ChAT) and tyrosine hydroxylase (TH) immunohistochemistry. In the dorsal tegmentum, ChAT immunoreactive neurons were distributed in the parabrachial area [the pedunculopontine group (PPG)] and along the medial adjacent central gray [the lateral dorsal tegmental group (LDT)]. The cholinergic neurons in the LDT area were larger than those in the PPG. When adjacent tissue sections were labeled with TH we noted extensive overlap between catecholamine and cholinergic neurons in the PPG, suggesting that REM sleep may occur as a result of an interaction between these transmitters in this area rather than the medial pontine reticular formation where no cholinergic or catecholamine neurons were found. Cholinergic neurons were also found in the cranial nerve nuclei and the nucleus ambiguus. The presence of cholinergic neurons in the PPG and LDT suggest that these neurons may play an important role in the generation of some of the tonic and phasic components of REM sleep, such as cortical desynchronization, pontogeniculo occipital waves, and muscle atonia.     

Tonic and phasic respiratory drives to human genioglossus motoneurons during breathing

A tongue muscle, the genioglossus (GG), is important in maintaining pharyngeal airway patency. Previous recordings of multiunit electromyogram (EMG) suggest it is activated during inspiration in humans with some tonic activity in expiration. We recorded from populations of single motor units in GG in seven subjects during quiet breathing when awake. Ultrasonography assisted electrode placement. The activity of single units was separated into six classes based on a step-wise analysis of the discharge pattern. Phasic and tonic activities were analyzed statistically with the coefficient of determination (r2) between discharge frequency and lung volume. Of the 110 motor units, 29% discharged tonically without phasic respiratory modulation (firing rate approximately 19 Hz). Further, 16% of units increased their discharge during expiration (expiratory phasic and expiratory tonic units). Only half the units increased their discharge during inspiration (inspiratory phasic and inspiratory tonic units). Units firing tonically with an inspiratory increase had significantly higher discharge rates than those units that only fired phasically (peak rates 25 vs. 16 Hz, respectively). Simultaneous recordings of two or three motor units showed neighboring units with differing respiratory and tonic drives. Our results provide a classification and the first quantitative measures of human GG motor-unit behavior and suggest this activity results from a complex interaction of inspiratory, expiratory, and tonic drives at the hypoglossal motor nucleus. The presence of different drives to GG implies that complex premotor networks can differentially engage human hypoglossal motoneurons during respiration. This is unlike the ordered recruitment of motor units in limb and axial muscles.     

Sleep-related increase in activity of mesopontine neurons in old rats

Relationships between age-related changes in sleep patterns and neuronal activity have received scant attention. In the present study, reticularis pontis oralis (RPO) and ventral tegmental nucleus of Gudden (VTN) neurons were recorded in unanesthetized restrained young (3 months) and old (23 months) Sprague-Dawley rats during wakefulness (W), slow wave sleep (SWS) and rapid eye movement (REM) sleep. All RPO neurons displayed a tonic activity. Firing rates were similar during W in young and old rats. In contrast, firing rates were higher during SWS in old rats (P < 0.001). In both young and old rats, firing rates increased significantly during REM sleep as compared to W and SWS but this increase was markedly greater in old rats. Neurons recorded from VTN displayed bursting activity at theta frequencies during W and REM sleep. The frequency of VTN bursting neurons was higher during REM sleep as compared to W in both groups of age. This difference was significantly more pronounced in old as compared to young rats (P < 0.001). Sleep-related hyperactivity of pontine neurons is discussed in terms of a possible deficit in inhibitory processes in old rats.