Inspiratory on-switch evoked by mesencephalic stimulation: activity of medullary respiratory neurones

Summary The activity of medullary inspiratory and expiratory neurones was studied in urethan-chloralose anaesthetized cats during stimulus — evoked inspiratory phase (inspiratory on-switch). All neurones were characterized according to their axonal destination (i.e. bulbospinal neurones or vagal motoneurones) or the absence of such axonal projections (i.e. propriobulbar neurones), and to their location in the dorsal or ventral respiratory nuclei. 1. The inspiratory on-switch effects were elicited during expiration (E phase) by brief tetanic electrical stimulation (50 to 100 ms duration; 0.5 mA; 300 Hz) delivered to the mesencephalic periaqueductal central gray and the adjacent reticular formation. The evoked inspiratory effects observed on the phrenic nerve discharge consisted of: (i) an immediate response (latency 20 ± 5 ms) of stable duration related to the stimulus (primary response: Prim.R.), (ii) a delayed response (patterned response: Patt.R.) appearing after a latent period (silent phase: Sil.P.) of 100 ms maximal duration. The later the stimulus in the E phase, the longer was the duration of the Patt.R. (300 to 1000 ms). 2. The stimulation evoked an earlier activation of the inspiratory bulbospinal neurones (latency 12 ± 6 ms) than that obtained in the phrenic nerve (Prim.R.). Hence, the Prim.R. originated from the bulbospinal pathway and not from a pathway directly impinging on the motoneurones. Conversely during stimulation very few inspiratory propriobulbar neurones were activated and no expiratory neurone discharged. 3. During the phrenic Sil.P., 46% of the inspiratory bulbospinal neurones continued to discharge with a firing rate lower than that during the stimulus train, while most of the inspiratory propriobulbar and expiratory neurones were not active. 4. During the Patt.R. all inspiratory bulbospinal neurones discharged early and were strongly activated whatever the Patt.R. duration whereas the expiratory neurones were not active. Inspiratory propriobulbar neurones were either not recruited or recruited later, and the number of active neurones increased as the duration of the Patt.R. lengthened. 5. Our results suggest that the eliciting of the stimulus-evoked inspiration (Patt.R.) primarily depends on the activation of the inspiratory bulbospinal neurones. These neurones therefore would not only be the output neurones of the medullary respiratory centres, but they would serve other roles such as building up of the excitation in other respiratory neurones, thus acting as a component of the inspiratory ramp generator.Abbreviations Prim.R Primary response - Patt.R Patterned response - Sil.P Silent phase - I phase Inspiratory phase - E phase Expiratory phase - IBSN Inspiratory bulbospinal neurones - IPBN Inspiratory propriobulbar neurones - EBSN Expiratory bulbospinal neurones - EPBN Expiratory propriobulbar neurones - DRN Dorsal respiratory nucleus - VRN Ventral respiratory nucleus Supported by CNRS (LA 205 and ATP no 4188) and Fondation pour Ia recherche médicale     

The mode of synaptic activation of pyramidal neurons in the cat primary somatosensory cortex: an intracellular HRP study

Summary A total of 141 pyramidal neurons in the cat primary somatosensory cortex (SI) were recorded intracellularly under Nembutal anesthesia (7 in layer II, 43 in layer III, 8 in layer IV, 58 in layer V and 25 in layer VI). Most neurons were identified by intracellular staining with HRP, though some layer V pyramidal neurons were identified only electrophysiologically with antidromic activation of medullary pyramid (PT) or pontine nuclear (PN) stimulation. Excitatory synaptic potentials (EPSPs) were analyzed with stimulation of the superficial radial nerve (SR), the ventral posterolateral nucleus (VPL) in the thalamus and the thalamic radiation (WM). The pyramidal neurons in layers III and IV received EPSPs at the shortest latency: 9.1±2.1 ms (Mean+S.D.) for SR and 1.6±0.7 ms for VPL stimulation. Layer II pyramidal neurons also responded at a short latency to VPL stimulation (1.7±0.5 ms), though their mean latencies for SR-induced EPSPs were relatively longer (10.6±1.9 ms). The mean latencies were much longer in layers V and VI pyramidal neurons (10.2±2.4 ms and 2.9±1.5 ms in layer V pyramidal neurons and 9.9±2.5 ms and 2.8±1.6 ms in layer VI pyramidal ones, respectively for SR and VPL stimulation). The comparison of the latencies between VPL and WM stimulation indicates that most layer III–IV pyramidal neurons and some pyramidal cells in layers II, V and VI received monosynaptic inputs from VPL. These findings are consistent with morphological data on the laminar distribution of thalamocortical fibers, i.e., thalamocortical fibers terminate mainly in the deeper part of layers III and IV with some collaterals in layers V, VI and II-I. The time-sequences of the latencies of VPL-EPSPs indicate that corticocortical and/or transcallosal neurons (pyramidal neurons in layers II and III) fire first and are followed by firing of the output neurons projecting to the subcortical structures (pyramidal neurons in layers V and VI).     

Integration in descending motor pathways controlling the forelimb in the cat. 13. Corticospinal effects in shoulder,elbow, wrist,and digit motoneurones

Summary The effect of corticospinal volleys evoked by stimulation of the contralateral pyramid was investigated using intracellular recordings from motoneurones to forelimb muscles. Confirming and extending previous observations (Illert et al. 1977, lllert and Wiedemann 1984), short latency EPSPs within a disynaptic range were evoked by a train of pyramidal volleys in all varieties of shoulder, elbow, wrist and digit motoneurones. The amplitude of pyramidal EPSPs was sensitive to the stimulus repetition rate. Maximal amplitudes were observed around 2–4 Hz, while at 10 Hz the early EPSP was markedly reduced and the long latency EPSP abolished. The persistence of disynaptic EPSPs after a corticospinal transection in C5/C6 suggested that, for all types of forelimb motor nuclei, disynaptic EPSPs are relayed by C3–C4 propiospinal neurones (PNs) (c.f. Illert et al. 1977). The transection, however, caused a clear reduction in the EPSP of all motoneurone types. After a ventral lesion of the lateral funicle in C5/C6 interrupting the axons of the C3–C4 PNs, disynaptic (and possibly trisynaptic) EPSPs were evoked by a short train of pyramidal volleys. It is postulated that intercalated neurones in a disynaptic cortico-motoneuronal pathway also exist in the forelimb segments. Disynaptic pyramidal IPSPs were observed in most types of forelimb motor nuclei both before and after a corticospinal transection in C5/C6. At all joints, pyramidal excitation dominated in motoneurones to physiological flexors, while in extensor motoneurones mixed excitation and inhibition or dominant inhibition was common. Comparison of pyramidal effects in slow motoneurones (classified according to the after-hyperpolarization duration) to the long head of the triceps and anconeus revealed dominant excitation in the former and inhibition in the latter. It is suggested that the slow motor units in these muscles differ in their function although both muscles are elbow extensors.This work was supported by the Swedish Medical Research Council (project no. 94 and 6953)     

Neuronal relays in double crossed pathways between feline motor cortex and ipsilateral hindlimb motoneurones

Coupling between pyramidal tract (PT) neurones and ipsilateral hindlimb motoneurones was investigated by recording from commissural interneurones interposed between them. Near maximal stimulation of either the left or right PT induced short latency EPSPs in more than 80% of 20 commissural interneurones that were monosynaptically excited by reticulospinal tract fibres in the medial longitudinal fascicle (MLF). The EPSPs were evoked at latencies that were only 1–2 ms longer than those of EPSPs evoked from the MLF, compatible with a disynaptic coupling between PT fibres and these commissural interneurones. EPSPs evoked by PT stimulation were frequently associated with IPSPs which either followed or preceded the EPSPs. The latencies of the IPSPs (on average about 1 ms longer than latencies of the earliest EPSPs) indicated that they were mediated via single additional inhibitory interneurones. Records from a sample of nine commissural interneurones from a different population (with monosynaptic input from group I and/or II muscle afferents, and disynaptically excited from the MLF) suggest that actions of PT fibres on such interneurones are weaker because only four of them were excited by PT stimuli and at longer latencies. By demonstrating disynaptic coupling between PT neurones and commissural interneurones via reticulospinal fibres, the results provide a direct demonstration of trisynaptic coupling in the most direct pathways between PT neurones and ipsilateral motoneurones, and thereby strengthen the proposal that the double crossed pathways between PT neurones and ipsilateral motoneurones might be used to replace crossed actions of damaged PT neurones.     

Electrical stimulation of arterial and central chemosensory afferents at different times in the respiratory cycle of the cat: II. Responses of respiratory muscles and their motor nerves

The response patterns of the electrical activity of the respiratory motor nerves and muscles to brief electrical stimulation of the arterial and the intracranial chemosensory afferents were studied in anesthetized cats. Stimulation during inspiration increased the activity of phrenic nerve and the inspiratory muscles (intercostal, diaphragm) with a latency of 15–25 ms, whereas expiratory muscle activity in the following expiration remained almost unaltered. Stimulation during expiration increased the activity of expiratory nerves and muscles (intercostal, abdominal) after a delay of 80–120 ms. The later the stimulation occurred in the insor expiratory period the larger the increase in amplitude and in steepness of rise of the respective integrated activity in respiratory nerves and muscles. Stimulation in early inspiration shortened the discharge period of inspiratory muscles, whereas excitation in early expiration caused an earlier onset and prolonged the activity in the expiratory muscles. Stimulation in the late phase of ins- or expiration prolonged the discharge of the respective nerves and muscles. Both the arterial (carotid sinus nerve, CSN, and aortic nerve, AN) and intracranial chemosensory (VM) afferents stimuli were able to affect both the inspiratory and the expiratory mechanisms. The restriction of the effects to the phase of the stimulus suggests a mechanism by which these afferents, when activated during inspiration, effectively project only to inspiratory neurones, and vice versa for expiration.Supported by the Deutsche Forschungsgemeinschaft, SFB 114 Bionach     

Comparison of human motor cortical projections to abdominal muscles and intrinsic muscles of the hand

Summary Percutaneous electrical stimulation of the motor cortex was used to activate rapidly conducting corticofugal pathways to human abdominal muscles. Following cortical stimulation the response latencies for the abdominal muscles were similar to those for limb muscles which are a similar distance from the motor cortex. Cortically evoked responses recorded from the abdominal muscles had the same latency and similar amplitude during several voluntary tasks including expiration, expulsive manoeuvres and trunk flexion. Responses could also be evoked when the chemical drive to breathe was increased by rebreathing. In addition, the properties of the cortical projection to muscles of the abdominal wall were directly compared with those of the projection to the intrinsic muscles of the hand. The latencies of responses in abdominal muscles and intrinsic muscles of the hand were measured during static contractions over a range of strengths in the same subjects (0–100% maximal voluntary contraction, MVC). For both muscle groups, cortically evoked muscle responses of minimal latency occurred when background contractions reached 10–20% MVC with responses of maximal amplitude at 60% MVC. The variability in latency of fifty consecutive responses were similar for the two muscle groups. Furthermore, post-stimulus time histograms for 4 rectus abdominis motoneurones revealed a brief initial excitatory peak of 1.15ms duration (range 0.96–1.34ms) following cortical stimulation. The characteristics of this peak are the same as reported for motoneurones of intrinsic hand muscles. These findings demonstrate a powerful rapidly conducting pathway from the motor cortex to the human abdominal muscles. This pathway has many of the same properties as the monosynaptic corticospinal projection to the distal muscles of the upper limb.     

Convergence of skin reflex and corticospinal effects in segmental and propriospinal pathways to forelimb motoneurones in the cat

The organization of facilitatory convergence from cutaneous afferents (Skin) and the corticospinal tract (pyramidal tract, Pyr) in pathways to forelimb motoneurones of mainly distal muscles was studied in anaesthetized cats by analysing postsynaptic potentials (PSPs), which were spatially facilitated by combinations of stimuli to the two sources at different time intervals. Conditioning Pyr volleys facilitated Skin-evoked PSPs of fixed (1.2–3.6 ms) central latencies (Skin PSPs), suggesting that disynaptic and polysynaptic skin reflex pathways are facilitated from the pyramidal tract. The shortest latencies (1.2–1.7 ms) of pyramidal facilitation suggested direct connection of pyramidal fibres with last order neurones of skin reflex pathways. Conditioning Skin volleys facilitated Pyr-evoked PSPs of fixed, mostly disynaptic latencies (1.0–2.5 ms; Pyr PSPs), suggesting that pyramido-motoneuronal pathways are facilitated from Skin at a premotoneuronal level. The shortest pathway from skin afferents to the premotor neurones appeared to be monosynaptic. Although Pyr and Skin volleys were mutually facilitating, the facilitation curve of Pyr PSPs and that of Skin PSPs were discontinuous to each other, with the peak facilitation at different Skin-Pyr volley intervals. Transection of the dorsal column (DC) at the C5/C6 border had little effect on the latencies or amplitudes evoked by maximal stimulation and the pyramidal facilitation of Skin PSPs. In contrast, the facilitation of Pyr PSPs by Skin stimulation was greatly decreased after the DC transection, and the facilitation curve of Pyr PSPs was continuous to that of Skin PSPs, with no separate peak. Latencies of Pyr PSPs ranged similarly to those in DC intact preparations. More rostral DC transection (C4/C5 border) reduced Skin-facilitated Pyr excitatory PSPs (EPSPs) less than C5/C6 lesions, suggesting that the C5 segment also contains neurones mediating Skin-facilitated Pyr EPSPs. The results show that convergence from skin afferents and the corticospinal tract occurs at premotor pathways of different cervical segments. We suggest that corticospinal facilitation of skin reflex occurs mostly in the brachial segments and Skin facilitation of cortico-motoneuronal effects takes place largely in the rostral cervical segments and partly in the brachial segments.     

Pyramidal excitation in long propriospinal neurones in the cervical segments of the cat

Summary 1. The effect of stimulating the contralateral pyramid has been investigated with intracellular recording from 128 long propriospinal neurones (long PNs) in the C3-Th1 segments of the cat. Long PNs were identified by the antidromic activation from the Th13 segment. They were located in laminae VII–VIII of Rexed. Single pyramidal stimulation evoked monosynaptic EPSPs in 15/40 of the long PNs in cats with intact pyramid. In 15 other long PNs, a train of three to four pyramidal stimuli evoked EPSPs with latencies indicating a minimal disynaptic linkage. The remaining 25% of the long PNs lacked mono- or disynaptic pyramidal EPSPs. In a few cases longer latency excitation was observed. 2. The location of the intercalated neurones which mediate the disynaptic pyramidal EPSPs was investigated by making four different lesions of the corticofugal fibres: 1) at the border of the C5 and C6 segments, 2) at the border of the C2 and C3 segments, 3) at the caudal part of the pyramid; three mm rostral to the decussation and 4) at the level of the trapezoid body. Stimulation of the corticofugal fibres was made either rostral to lesion 3 (rPyr) in order to activate neurones in a cortico-bulbospinal pathway or caudal to lesion 3 (cPyr) to activate neurones in a corticospinal pathway. In the former case, in one experiment, stimulation was made in the pyramid between lesions 3 and 4 (double pyramidal lesion). In case of cPyr stimulation, lesions 1 and 2 were added sequentially in order to investigate if the corticospinal excitation was mediated via C3–C4 PNs. All lesions were made mechanically, except lesion 2 which in some of the experiments was performed by reversible cooling. 3. Stimulation in the pyramid rostral to lesion 3 and in between lesions 3 and 4 evoked disynaptic EPSPs in the long PNs, which shows that they were mediated via reticulospinal neurones. Stimulation in cPyr after lesion 3 elicited disynaptic EPSPs, which remained after lesion 1 but were abolished after adding lesion 2. It is concluded that the disynaptic cPyr EPSPs were mediated via intercalated neurones in the C3–C4 segments. 4. When the disynaptic cPyr EPSP was conditioned with a single volley in nucleus ruber and/or in tectum, it was markedly facilitated, especially when the conditioned volley was applied simultaneously with the effective cPyr volley. The results show that the intercalated neurones in the C3–C4 segments receive monosynaptic convergence from cortico-, rubro- and tectospinal] fibres. Stimulation in the lateral reticular nucleus (LRN) evoked monosynaptic EPSPs. These EPSPs had similar latencies and shapes as those previously recorded in forelimb motoneurones and which have been shown to be due to activation of ascending branches of the C3–C4 PNs. This finding in addition to the striking similarity of the descending input pattern of long PNs as compared to the forelimb motoneurones strongly suggest that short C3–C4 PNs project both to long PNs as well as to forelimb motoneurones. 5. Spatial facilitation of disynaptic EPSPs in long PNs was also observed between rPyr volleys and tectal volleys. The results suggest that common reticulospinal neurones which project to the long PNs receive monosynaptic convergence from corticofugal and tectofugal fibres but in some of the reticulospinal neurones the main input is cortical and in others tectal. Monosynaptic EPSPs were evoked from the medial part of the reticular formation, from 2 mm caudal to 6 mm rostral of the obex level. These EPSPs were presumably due to direct activation of reticulospinal neurones. 6. Convergence of disynaptic excitation mediated by cortico-propriospinal and cortico-reticulospinal routes was observed in about 12% of the long PNs. Convergence of monosynaptic corticospinal and disynaptic corticoreticulospinal and/or cortico-propriospinal input was observed in about 15% of the long PNs. 7. The role of the monosynaptic pyramidal input and disynaptic corticoreticulospinal and cortico-propriospinal (mediated by short C3–C4 PNs) inputs to long PNs is discussed in relation to postural control during movements of head and forelimb.     

Integration in descending motor pathways controlling the forelimb in the cat. 1. Pyramidal effects on motoneurones

Summary Stimulation of the contralateral pyramid and intracellular recording from forelimb motoneurones was used to investigate corticomotoneuronal pathways in the cat.A train of pyramidal volleys evokes short-latency EPSPs in flexor motoneurones and in many extensor motoneurones. The latency for the on-set after the effective pyramidal volley — usually the third — strongly indicates a disynaptic linkage. These disynaptic EPSPs were common in triceps motoneurones to fast heads but rare in those to slow heads.Pyramidal IPSPs with a slightly longer latency, suggesting a trisynaptic linkage, were found in both flexor and extensor motoneurones. They were common in motoneurones to slow heads of triceps. Disynaptic pyramidal IPSPs were found only occasionally.In addition pyramidal volleys may evoke late large EPSPs and/or IPSPs in any combination with the short-latency PSPs.Supported by the Deutsche ForschungsgemeinschaftIBRO/UNESCO Fellow     

Trigeminal excitation of dorsal neck motoneurones in the cat

Summary Excitation of dorsal neck motoneurones evoked by electrical stimulation of primary trigeminal afferents in the Gasserian ganglion has been investigated with intracellular recording from -motoneurones in the cat. Single stimulation in the Gasserian ganglion ipsi-and contralateral to the recording side evoked excitatory postsynaptic potentials (EPSPs) in motoneurones innervating the lateral head flexor muscle splenius (SPL) and the head elevator muscles biventer cervicis and complexus (BCC). The gasserian EPSPs were composed of early and late components which gave the EPSPs a hump-like shape. A short train of stimuli, consisting of two to three volleys, evoked temporal facilitation of both the early and late EPSP components. The latencies of the gasserian EPSPs ranged from 1.6 to 3.6 ms in SPL motoneurones and from 1.6 to 5.8 ms among BCC motoneurones. A rather similar latency distribution between 1.6 and 2.4 ms was found for ipsi- and contralateral EPSPs in SPL and BCC motoneurones, which is compatible with a minimal disynaptic linkage between primary trigeminal afferents and neck motoneurones. Systematic transections of the ipsi- and contralateral trigeminal tracts were performed in the brain stem between 3 and 12 mm rostral to the level of obex. The results demonstrate that both the ipsi- and contralateral disynaptic and late gasserian EPSPs can be mediated via trigeminospinal neurones which take their origin in the nucleus trigeminalis spinalis oralis. Transection of the midline showed that the contralateral trigeminospinal neurones cross in the brain stem. Systematic tracking in and around the ipsilateral trigeminal nuclei demonstrated that the axons of ipsilateral trigeminospinal neurones descend just medial to and/or in the medial part of the nucleus. Spinal cord lesions revealed a location of the axons of the ipsilateral trigeminospinal neurones in the lateral and ventral funiculi. Interaction between the ipsi- and contralateral gasserian EPSPs showed complete summation of the disynaptic EPSP component, while the late components were occluded by about 45%. These results show that the disynaptic EPSPs are mediated by separate trigeminospinal neurones from the ipsi- and contralateral side, while about half of the late EPSPs are mediated by common neurones which receive strong bilateral excitation from commissural neurones in the trigeminal nuclei. Spatial facilitation was found in the late gasserian EPSP but not in the disynaptic gasserian EPSP by conditioning stimulation of cortico- and tectofugal fibres. Disynaptic pyramidal and tectal EPSPs, which are mediated by reticulospinal neurones, were facilitated by a single stimulation in the gasserian ganglion at an optimal interval of 2 ms. It is suggested that primary trigeminal afferents can excite the reticulospinal neurones via a disynaptic trigeminoreticular pathway.     

Monosynaptic excitatory connexions of reticulospinal neurones in the nucleus reticularis pontis caudalis with dorsal neck motoneurones in the cat

Summary 1. Projections of reticulospinal neurones (RSNs) in the nucleus reticularis pontis caudalis (N.r.p.c.) to dorsal neck motoneurones supplying splenius (SPL, lateral head flexor) and biventer cervicis and complexus (BCC, head elevator) muscles were studied in the cat anaesthetized with pentobarbiturate or -chloralose. 2. Threshold mapping for evoking antidromic spikes revealed that most of RSNs tested projecting down to brachial segments but not to lumbar segments (C-RSNs) gave off collaterals to the gray matter of the upper spinal cord in C2–C3 segments. 3. Spike triggered averaging showed that negative field potentials were evoked after firing of a single C-RSN (single fibre focal synaptic potentials, FSPs) in the region of C2–C3 where large antidromic field potentials from nerves supplying SPL or BCC muscles were evoked. The single fibre FSPs ranged between 1 and 10 V in amplitude and had latencies between 0.7 and 1.2 ms from the onset of the triggering spike. In most cases, a presynaptic spike preceded the negative potential by 0.3 ms. These results indicated that C-RSNs project to the SPL or BCC motor nucleus. 4. Spike triggered averaging of postsynaptic potentials revealed EPSPs (single fibre EPSPs) in 36 dorsal neck motoneurones, predominantly in SPL (25) and less in BCC (11) motoneurones, evoked from 15 C-RSNs. The amplitude of the single fibre EPSPs ranged from 5 to 310 V, and had latencies of 0.8–2.0 ms from the onset of the triggering spikes of C-RSNs, or 0.3–0.5 ms from the presynaptic spike when recorded. The results indicated monosynaptic excitatory connexions of C-RSNs to dorsal neck motoneurones. 5. Single fibre EPSPs from a C-RSN were usually recorded from either BCC or SPL motoneurones but not from both types of motoneurones, when tested in many motoneurones. This showed that connexions of C-RSNs with dorsal neck motoneurones were muscle specific. 6. RSNs projecting down to the lumbar segment (L-RSN) also showed branching in C2–C3 segments. Excitatory monosynaptic connexion of L-RSNs with neck motoneurones were demonstrated by recording single fibre postsynaptic population potentials (p.s.p.p.s.) from the C2 ventral root perfused with sucrose. The probability of evoking monosynaptic single fibre p.s.p.p.s. was less (19%) than for C-RSNs (59%).     

The Origin of a Descending Pathway with Monosynaptic Action on Flexor Motoneurones

The postsynaptic effects evoked in lumbar motoneurones were studied following electrical stimulation of the brain stem in the cat. The spinal cord was transected at the lower thoracic level leaving only the ipsilateral ventral quadrant intact. With a stereotactical method a low threshold focus was found in the medial brain stem from which monosynaptic EPSPs could be evoked in flexor motoneurones. It is concluded that this effect is mediated by fibres descending in the ipsilateral medial longitudinal fascicle and it is tentatively suggested that these fibres originate from the ipsilateral upper medullary or lower pontine reticular formation. Monosynaptic EPSPs were also evoked in some extensor motoneurones from this medial brain stem region and at such a strength of stimulation that stimulus escape to the lateral vestibulospinal tract is excluded.     

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

Summary 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 averarging 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 moteneurones. 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). Eikewise, 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-, diand/or polysynaptically and are mainly concerned with the control of vertical head movements.     

The vibrissal pad as a source of sensory information for the oculomotor system of the cat

Summary Responses from lateral rectus, medial rectus and retractor bulbi nerves were obtained following electrical stimulation of the vibrissal pad of the cat. Discharges in afferent fibres dissected from the infraorbital nerve were recorded during movements of the vibrissae and following electrical stimulation of the vibrissal pad. Both stimuli activated the same population of A fibers. Intracellular records were obtained from lateral rectus motoneurones identified antidromically in the principal abducens nucleus and from retractor bulbi motoneurones similarly identified in the accessory abducens nucleus. EPSPs (3 mV) were recorded in lateral rectus motoneurones following electrical stimulation of the ipsilateral vibrissal pad at a latency of 3.5 ms. Large-amplitude disynaptic EPSPs (15 mV) were recorded in retractor bulbi motoneurones following the same vibrissal stimulation. The synaptic excitation evoked in both lateral rectus and retractor bulbi motoneurones through stimulation of the ipsilateral vibrissal pad induced an early retraction followed by an abduction of the eye ball. The hypothesis is that the vibrissal message might complement other sensory modalities in the generation of patterned eye movements.Supported by CNRS (GR 45) and by Grant DGRST No. 78.7.3017     

Serotonin reveals ineffective spinal pathways to contralateral phrenic motoneurons in spinally hemisected rats

Serotonin reveals ineffective (subthreshold) pathways from the C2 lateral funiculus to ipsilateral phrenic motoneurons in spinalized rats. The objective of the present study was to investigate serotonergic modulation of crossed-spinal pathways to contralateral phrenic motoneurons. Rats (n = 10) were anesthetized (urethane), paralyzed, vagotomized, and artificially ventilated. The spinal cord was hemisected at C1–C2 and, on the intact side, a tungsten stimulating electrode was placed ventral to the C2 dorsal root entry zone in the dorsolateral ( 1.1 mm) or the ventrolateral funiculus (2.2 mm depth). Single shocks (100–750 A, 0.1–0.5 ms, 2 Hz) elicited a short-latency ( 1.0 ms to peak) excitation in the ipsilateral phrenic nerve, but usually evoked little or no response in the contralateral phrenic nerve at either stimulus site. Following systemic injection of the monoamine oxidase inhibitor pargyline (25 mg/kg) and the serotonin precursor 5-hydroxytryptophan (5–10 mg/kg), complex responses were revealed in the contralateral phrenic nerve, including; (1) spontaneous tonic activity; (2) a short-latency (1.0 ms to peak) evoked excitation; and (3) two long-latency (2.2 and 7.8 ms to peak) evoked excitations. The longest latency excitation was expressed only when the stimulating electrode was positioned in the dorsolateral funiculus. Contralateral evoked responses were blocked by systemic methysergide (2–6 mg/kg), a broad-spectrum serotonin receptor antagonist. These results indicate that serotonin converts ineffective crossed phrenic pathways in the spinal cord to effective pathways. It remains to be determined whether serotonin is both necessary and sufficient in this modulatory process, or if it is a nonspecific result of increased phrenic motoneuron excitability.     

Integration in descending motor pathways controlling the forelimb in the cat 14. Differential projection to fast and slow motoneurones from excitatory C3-C4 propriospinal neurones

Summary The projection of C3-C4 propriospinal neurones (PNs) to -motoneurones of forelimb muscles has been analysed with the aid of antidromic stimulation of the ascending branch of the PNs to the lateral reticular nucleus (LRN). A single stimulus of 500 A applied in the caudo-dorsal part of the LRN evoked a maximal or > 90% maximal monosynaptic EPSP in the motoneurones. Systematic mapping of EPSPs evoked by stimulation of 500 A in and around the LRN revealed that at this strength there was hardly any co-activation of a medial system (Peterson et al. 1979) which evoked small monosynaptic EPSPs with shorter latency and faster time course. The LRN EPSP amplitude was positively correlated with the homonymous group Ia EPSP amplitude, the input resistance and the afterhyperpolarization (AHP) duration. It is therefore postulated that the LRN EPSP amplitude is correlated with motor unit type (Burke 1967, 1968; Burke et al. 1973) with the largest EPSPs in slow (S), the smallest in fast, fatiguable (FF) and possibly intermediate sized in fast, fatigue resistant (FR) units. There was only a small difference in latency of the LRN EPSP in fast and slow motoneurones, while the time course was considerably slower in the latter. It is suggested that slow motoneurones receive projection both from fast and slowly conducting PNs but fast motoneurones mainly from fast PNs. Comparison of the disynaptic pyramidal EPSPs and the LRN EPSPs revealed a positive correlation, but the amplitude ratio pyramidal EPSP: LRN EPSP was smaller in slow than in fast motoneurones. A negative correlation was found between this amplitude ratio and the latency of the disynaptic pyramidal EPSP. It is suggested that this correlation reflects the excitability level in the PNs and that low excitability is due to inhibition of the PNs.This work was supported by the Swedish Medical Research Council (project no. 94 and 6953)     

Medullary control of the pontine swallowing neurones in sheep

Summary The origin of the inputs from the medullary swallowing centre (dorsal region including the nucleus of the solitary tract, or ventral region corresponding to the reticular formation surrounding the nucleus ambigous) to the pontine swallowing neurones (PSNs) was studied in sheep anaesthetized with halothane.Out of 101 PSNs located in the posterior part of the trigeminal (Vth) motor nucleus, 46 were activated by stimulating either the dorsal (21 neurones) or the ventral (25 neurones) region of the ipsilateral medullary swallowing centre, 3–4 mm rostral from the obex. Thirty-one neurones out of the 46 were identified as a motoneurones supplying swallowing muscles (mylohyoïd, anterior body of digastric and medial pterygoïd). Their average activation latency through stimulation of the dorsal medullary region was about 1 ms longer than through stimulation of the ventral region (3.63 ms±0.81 versus 2.72 ms±0.32).To determine the origin of the medullary input to the PSNs, we tried to activate the medullary swallowing neurones (MSNs) antidromically through stimulating the posterior part of the Vth motor nucleus, which contains the swallowing motoneurones. Seventy-three MSNs were tested (25 located in the dorsal and 48 in the ventral region). None of the dorsal neurones tested could be antidromically activated by pontine stimulation: 15 ventral neurones showed a clear antidromic response (collision test) with an average latency of 2.5 ms±0.73. These neurones, which send their axons into the pons, were all located in the reticular formation, above the nucleus ambiguus, 3–4 mm rostral from the obex.These results suggest that MSNs in the ventral reticular formation connect the medullary swallowing centre to the Vth motor nucleus. They also suggest that during swallowing, inputs originating from the dorsal region of the medullary centre (interneurones programming the motor sequence) are relayed in the ventral region (reticular formation adjacent to the nucleus ambiguus) before reaching the PSNs.This work was supported, in part, by grants from CNRS (LA 205), INRA and M.R.I. (82 E 0685)     

Influence of rubrospinal tract and the adjacent mesencephalic reticular formation on the activity of medullary respiratory neurons and the phrenic nerve discharge in the rabbit

Suprapontine brain sites acting on the central respiratory system have been demonstrated to give rise to inspiratory as well as expiratory facilitatory effects. In the present study the inspiratory inhibitory effect which has been reported in the cat to be elicited consistently by electrical stimulation of the rubrospinal tract and the adjacent mesencephalic reticular formation was examined in the urethane-anaesthetized rabbit. Stimulation of these sites with single electrical shocks of moderate intensity induced a short latency (onset after 3.0 ms) transient (duration: 29 ms) inhibition of the phrenic nerve activity (PHR). Short volleys of stimuli applied in mid- to late-inspiration led to a premature off-switch of inspiration. The extracellularly recorded discharge activity of the different types of medullary respiration-related units (RRU) reflected these alterations, accordingly. Axonal connections of RRU with mesencephalic structures were evaluated. Examination of orthodromic responses of medullary RRU to stimulation of this pathway revealed that most bulbospinal inspiratory neurons (10 out of 13) were paucisynaptically inhibited after short latency (at least 1.2 ms). The conduction time from bulbospinal inspiratory neurons to the recording site of PHR was 1.6 ms. Thus, a disynaptic pathway — including bulbospinal inspiratory neurons — is suggested inducing inspiratory inhibition 3.0 ms after single shock midbrain stimulation. This inhibition results in disfacilitation of phrenic motoneurons. The fact that extensive electrolytic lesions of the pneumotaxic center in rostral pons did not abolish the observed inspiratory inhibitions excludes these structures from being involved. A direct pathway from the red nucleus and the adjacent reticular formation to phrenic nuclei of the spinal cord, however, can not be excluded from being involved in the demonstrated inspiratory inhibition. The described effects may play a role in behavioral or voluntary control of respiration.     

Synaptic interactions between respiratory neurons during inspiratory on-switching evoked by vagal stimulation in decerebrate cats

To elucidate neuronal mechanisms underlying phase-switching from expiration to inspiration, or inspiratory on-switching (IonS), postsynaptic potentials (PSPs) of bulbar respiratory neurons together with phrenic nerve discharges were recorded during IonS evoked by vagal stimulation in decerebrate and vagotomized cats. A single shock stimulation of the vagus nerve applied at late-expiration developed an inspiratory discharge in the phrenic neurogram after a latency of 79+/-11 ms (n = 11). Preceding this evoked inspiratory discharge, a triphasic response was induced, consisting of an early silence (phase 1 silence), a transient burst discharge (phase 2 discharge) and a late pause (phase 3 pause). During phase 1 silence, IPSPs occurred in augmenting inspiratory (aug-I) and expiratory (E2) neurons, and EPSPs in postinspiratory (PI) neurons. During phase 2 discharge, EPSPs arose in aug-I neurons and IPSPs in PI and E2 neurons. These initial biphasic PSPs were comparable with those during inspiratory off-switching evoked by the same stimulation applied at late-inspiration. In both on- and off-switching, phase-transition in respiratory neuronal activities started to arise concomitantly with the phrenic phase 3 pause. These results suggest that vagal inputs initially produce a non-specific, biphasic response in bulbar respiratory neurons, which consecutively activates a more specific process connected to IonS.