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.     

Axonal trajectories of single Forel's field H neurones in the mesencephalon,pons and medulla oblongata in the cat

Summary We studied axonal trajectories of single Forel's field H (FFH) neurones (n= 19) in the mesencephalon, pons and medulla by systematic antidromic threshold mapping in cats and differentiated them into two major types. Type I neurones were characterized by projections to the oculomotor nucleus (IIIn) and type II neurones by lack of projections to the IIIn. 2. Type I neurones (11/19) were further classified into three subtypes by the lowest level of projections; type Ic (n = 3) which projected to the cervical cord and type Ib (n = 7) which terminated at the ponto-medullary level and type Ia (n = 1) at more rostral level. In the mesencephalon, stem axons passed just lateral to the IIIn and projected collaterals to the IIIn and the ventral part of the periaqueductal gray matter. In the lower brain stem, stem axons of type Ib and Ic neurones passed in the dorsal part of the reticular formation or in the medial longitudinal fasciculus and projected collaterals to the dorsal part of the nucleus reticularis pontis caudalis (NRPC) and the nucleus reticularis gigantocellularis (NRG) and the reticular formation underlying the nucleus prepositus hypoglossi (PH) and the raphe region. Projections to the superior colliculus were observed in two cases. 3. Type II neurones (8/19) were classified into 2 type IIb projecting to the ponto-medullary reticular formation and 6 type IIc projecting to the cervical spinal cord. In the mesencephalon, stem axons passed through a more lateral region than those of type I and projected collaterals to the mesencephalic reticular formation and the red nucleus. In the lower brain stem, the stem axons passed in the ventral part of the reticular formation corresponding to the central tegmental tract and projected collaterals to the ventral part of the NRPC and NRG. Projections to the interstitial nucleus of Cajal, the inferior olive and the reticular formation underlying the PH were also observed. 4. The dorsal and ventral location of, respectively, stem axons of type I and type II neurones in the lower brain stem was confirmed in a larger number of neurones in experiments with restricted mapping. 5. There was not much difference in location of cell bodies of type I (totally n = 50) and type II (n = 46) neurones. The proportion of spinal-projecting neurones were larger in type II (21/46, 46%) than in type I (7/50, 14%) neurones.     

Mono- and disynaptic excitatory inputs from the superior colliculus to vertical saccade-related neurons in the cat Forel's field H

Summary Excitatory inputs to neurons in the Forel's field H (FFH) related to visually induced vertical saccades from the ipsilateral superior colliculus (SC) were investigated in chronically prepared alert cats. By stimulation of the deep or intermediate layer of the SC, upward augmenting neurons (ANs) and one long-lead downward burst neuron (BN) were found to be activated monosynaptically, while medium-lead BNs were activated disynaptically. The monosynaptically activated neurons were not antidromically activated from the oculomotor nucleus, whereas disynaptically activated neurons were also activated antidromically from the inferior rectus subdivision of the nucleus. These results suggest that an excitatory input to the FFH from the SC for inducing vertical saccades of visual origin first reaches upward ANs and/or long-lead downward BNs in the FFH, which in turn drive medium-lead BNs in the same area synapsing with motoneurons related to vertical eye movements.Research fellow from the Department of Pathophysiology, Hebei Medical College, China     

Activity of neurons in Forel's field H during orienting head movements in alert head-free cats

Single unit activities were recorded in Forel's field H (FFH) at the mesodiencephalic junction during orienting head movements in two alert cats under headfree conditions. Recordings were made of 63 neurons of which 20 showed phasic firing that preceded the onset of head movements by 20–100 ms and was temporally related to the dynamic phase of the orienting head movement. Nineteen of these neurons showed a preference for upward movements, while the remaining neuron preferred downward movements. Activities during orienting movements in eight different directions (each separated by 45°) were systematically analyzed for 12 of the 19 upward-preferring neurons. The activities were broadly tuned; in most of the neurons, maximum activity was observed for direct upward movements (+90°), but significant activity was also observed for ipsilateral and contralateral oblique upward movements (+45° and +135°). In these cases, the increase in activity preceded the onset of the movement. Some increase in activity was also observed for ipsilateral and contralateral horizontal, oblique downward and downward movements. However, the increase in activity in the latter cases occurred simultaneously with or lagged behind the onset of the movement and was often preceded by a decrease in activity. The same pattern of directional tuning was observed in the EMG of the biventer cervicis muscle, a target of FFH neurons. The preferred directions of the 12 upward-preferring neurons were estimated by calculating the vector sum of the activity and were distributed between +68° and +108°. The same amount of activity was observed for ipsilateral and contralateral oblique upward movements, suggesting that FFH neurons on both sides of the brainstem are equally activated even during oblique orienting. Input from the ipsilateral superior colliculus was investigated in 18 neurons, all of which were orthodromically activated with a latency of 0.8–1.8 ms, suggestive of a mono- or disynaptic excitatory connection. Seven neurons were identified as descending projection neurons by antidromic activation from the ipsilateral medullary reticular formation. Repetitive microstimulation of unilateral FFH induced oblique upward head movements and an accompanying torsional component, while simultaneous bilateral stimulation at comparable stimulus strength induced purely upward head movements. These results strongly suggest that the vertical component of orienting head movements is encoded by equal bilateral activation of the FFH.     

Direct inhibitory projection of pontine omnipause neurons to burst neurons in the Forel's field H controlling vertical eye movement-related motoneurons in the cat

Summary This study examines the nature of the efferent projection of omnipause neurons (OPNs) in the midline pontine tegmentum to medium-lead burst neurons (BNs) in the Forel's field H (FFH), both of which exhibit activities related to vertical eye movements, using chronically prepared alert cats. Antidromic spikes of the BNs evoked by oculomotor nucleus stimulation were suppressed by shortly preceding (less than 5 ms) microstimulation within the OPN area including actual recording sites of OPNs. Many OPNs were antidromically activated by microstimulation at recording sites of the BNs. Furthermore, systematic tracking in and around the FFH with the stimulating microelectrode substantiated that the OPNs issued axonal branches within the BN area. These results suggest direct inhibitory projection of OPNs to the BNs.     

Mesencephalic projections to the first cervical segment in the cat

Mesencephalic neurons projecting to the upper cervical spinal cord were examined by mapping the distributions of labeled cells after injecting fluorescent tracers or wheat-germ agglutinin conjugated to horseradish peroxidase (WGA-HRP) into the C1 segment. Injections into the central or deep regions of the ventral horn produced retrograde labeling in cells of several mesencephalic regions. The majority of cells were found contralaterally in the superior colliculus and red nucleus, and ipsilaterally in and around the interstitial nucleus of Cajal (INC), in the cuneiform region, and in the fields of Forel. Smaller numbers of cells were located in the periaqueductal gray matter, nucleus annularis, and magnocellular nucleus of the posterior commissure. Dorsomedial injections in the ventral horn near the ventral commissure labeled only a subset of these projections, including cells in the mesencephalic reticular formation adjacent to the INC and in the nucleus annularis. Dorsolateral injections labeled some cells in the superior colliculus and were particularly effective at labeling cells in the red nucleus. These results suggest that at least ten different cell groups project to the ventral horn of the first cervical segment. Most, but not all, groups originate from regions implicated previously in the control of eye or head movements.     

Crossed reticulo-reticular projections in the medulla, pons and mesencephalon

Summary Injection of radioactive leucine in various regions of the brain stem reticular formation has revealed the presence of ample crossed reticulo-reticular connections in the cat. The terminal area for the crossed fibers are almost mirror images of the injected sites. The findings made is another example that hitherto unknown fiber connections can be demonstrated by axoplasmic protein tracing.     

Thermosensory defects after cervical spinal cord lesions in the cat

Summary The behavioural thermosensitivity of cat paws was examined before and/or after restricted uni- and/or bilateral lesions had been made in the spinal cord between the first and fifth cervical segments. Unilateral lesions of the lateral funiculus, which involved at least its whole width at the level of the central canal, reproducibly were found to interfere with the contralateral sensitivity for temperature increases and/or decreases. No corresponding thermosensory deficiencies were found after unilateral lesions involving the ventral spinal quadrant or the dorsal funiculus. Various bilateral and combined lesions were made, but no cat ever developed thermoanaesthesia. The bilateral lesions included bilateral transections of: the middle parts of the lateral funiculi, the dorsal halves of the lateral funiculi, the dorsal funiculi, and the ventral spinal half.Most of our knowledge about peripheral behavioural thermosensitivity after spinal cord injury is based on observations of human patients, especially after anterolateral chordotomies. The present finding of contralateral thermosensory deficiencies after lesions of the middle part of the lateral funiculus fits with some of the clinical reports. The present failure to cause thermoanaesthesia, on the other hand, is inconsistent with the theory of a single ascending spinal pathway for behavioural thermo-sensitivity, which has emanated mainly from the clinical observations.     

Mapping of monoamine neurones and fibres in the cat lower brainstem and spinal cord

Summary The localization of monoaminergic neurones in the medulla oblongata and the pons, and the distribution of catecholaminergic fibres in the spinal cord of the cat were investigated by means of formaldehyde-induced (FIF) or glyoxylic-acid-induced (GIF) fluorescence. Four groups of catecholamine (CA)-containing neurones were found in the following regions: (1) in the ventrolateral medulla oblongata within and adjacent to the lateral reticular nucleus, beginning slightly rostral to the medullo-spinal junction and extending rostrally to the cranial third of the inferior olive; (2) in the commissural, medial and lateral nucleus of the solitary tract; (3) cranial to the first group, closely adjacent to the facial nucleus and the superior olive; and (4) in the dorsolateral pons distributed to different nuclei, namely the nucleus coeruleus and subcoeruleus, the Koelliker-Fuse nucleus, and the medial and lateral parabrachial nuclei. The indoleamine (IA)-containing cell bodies were in general confined to the raphe nuclei, namely the nucleus raphe pallidus, nucleus raphe obscurus, nucleus raphe magnus, nucleus raphe pontis, nucleus raphe dorsalis and the central superior nucleus. A few IA-neurones were located more laterally, especially dorsal and lateral of the cranial half of the inferior olive, around the root of the hypoglossal nerve, in the lateral tegmental field and the pontine central gray. In the spinal cord most CA-fibres were found in the intermediolateral cell column. Another dense accumulation of CA-fibres was located dorsally and laterally of the central canal. The ventral and dorsal horns also contained CA-nervefibres which were slightly more numerous in the sacral spinal cord than in the more rostral parts of the spinal cord.     

Descending neural projections to the spinal cord in the channel catfish,Ictalurus Punctatus

Retrograde transport of horseradish peroxidase was used to determine the descending projections to the spinal cord in an otophysan fish, the channel catfish, Ictalurus punctatus. The majority of cells projecting to the spinal cord are located in the reticular formation, which is organized into rhombomeric segments. Vestibulospinal neurons are located in the descending, magnocellular, and tangential octaval nuclei, as well as in the medial octavolateralis nucleus of the lateral line system. Cells in the facial lobe project to the spinal cord. Additionally, axons of cells of the trigeminal system and the nucleus of the lateral lemniscus project caudally into the spinal cord. In the midbrain, descending spinal projections arise from cells of the medial longitudinal fasciculus and the red nucleus. More rostrally, cells of the ventrolateral thalamus, dorsal periventricular hypothalamus, central pretectal and magnocellular preoptic nuclei also project to the cord. The results of this study indicate that there are a number of homologies in the descending systems of bony fishes and other vertebrate taxa, including tetrapods. We also provide further evidence that a red nucleus is present in the brains of bony fishes and is therefore a primitive vertebrate character antedating the evolution of tetrapods. Anat. Rec. 252:235–253, 1998. © 1998 Wiley-Liss, Inc.     

Spino-olivary projections from the upper cervical spinal cord: An experimental study using autoradiography and horseradish peroxidase

Summary Spino-olivary projections from segments C1 and C2 were examined in 17 cats using autoradiographic methods and in nine cats using the method of retrograde horseradish peroxidase (HRP) transport. Injections of ³H-leucine at the junction of the C1–C2 segments produced anterograde terminal labelling in two regions of the contralateral inferior olive, one in the rostromedial half of the dorsal accessory olive (DAO), the other in the caudal half of the medial accessory olive (MAO). Projections to the rostromedial DAO were best demonstrated when tracer labelled the ventromedial part of the dorsal horn, while projections to the caudal MAO were strongly labelled by injections in both the lateral and medial parts of the intermediate grey matter. Injections of HRP into the region of the inferior olive led to retrograde marking of cells in both regions of the contralateral spinal cord implicated by autoradiographic studies to have spino-olivary projections. Dense groupings of small rounded or fusiform cells were labelled contralaterally on the medial aspect of the dorsal horn in C1 and C2, while medium-sized multipolar cells were more sparsely distributed throughout intermediate laminae of C1-C5.Olivary projections from dorsal column nuclei were also examined and compared to those of spinoolivary projections. Injections of ³H-leucine into n. gracilis and cuneatus led to terminal labelling in three olivary regions, including the rostral DAO, the caudo-lateral DAO and the caudal MAO. Projections from the DCN to the rostral DAO and the caudal MAO overlapped with regions of projection from upper cervical segments although the territories occupied by DCN and upper cervical projections were not identical. Amino acid injections which were confined to n. cuneatus gave rise to terminal labelling in only the rostromedial DAO.Supported by a grant to the Medical Research Council Group in Neuroscience, University of Montreal. F.J.R. Richmond is funded by a Scholarship from the MRC. Funds for travel were provided by the Advisory Research Committee of Queen's UniversityPresent address: Playfair Neurosciences Unit, The Toronto Western Hospital, 399 Bathurst Street, Toronto, Ontario, Canada M5T 2S8     

The fiber projections from the dentate nucleus to the reticular formation of the brain stem in the rabbit

Summary The projections from the dentate nucleus to the reticular formation of the brain stem in rabbit have been examined by means of the Fink-Heimer technique. The fibers arising from the dentate nucleus primarily project to the reticular formation via the contralateral and ipsilateral descending limbs of the brachium conjunctivum. The descending fibers project bilaterally to the parvocellular reticular nucleus, the ventral reticular nucleus, the gigantocellular reticular nucleus, the nucleus raphe magnus, the oral and caudal pontine reticular nuclei, and the reticulo-tegmental nucleus.     

Descending projections to the inferior olive from the mesencephalon and superior colliculus in the cat

Summary Descending projections from the mesencephalon and superior colliculus to the inferior olive were analyzed by an autoradiographic tracing method. Injections of tritium-labelled leucine were placed in regions which had previously been identified as sources of afferents to the olive. These were located adjacent to the central gray and extended from the rostral red nucleus to the posterior thalamus. Additional injections were made in the superior colliculus. Other injections were placed in the basal ganglia and thalamus. Injections restricted to one side of the central mesencephalon resulted in predominantly ipsilateral labelling of the olive. After injections in the caudo-medial parafascicular and subparafascicular nuclei and rostral nucleus of Darkschewitsch, deposits of grains were observed in the rostral pole of the medial accessory olive and adjacent ventral lamella of the principal olive. The medial accessory olive contained grains into its middle third. More caudal injections which involved the interstitial nucleus of Cajal as well as the nucleus of Darkschewitsch and rostral red nucleus resulted in the dense labelling of the entire principal olive (except the dorsal cap), the entire medial acessory olive (except subnucleus and the caudo-medial pole) and the caudo-dorsal accessory olive. Injections centered in the caudal magnocellular red nucleus and extending into the rostral parvocellular division labelled the dorsal lamella of the principal olive almost exclusively. When only the caudal part of the red nucleus was involved in the injection, the olive was entirely clear of grains. Minor contralateral distributions were observed in the dorsomedial cell column, the medial tip of the dorsal lamella and in the caudal medial accessory olive. The deep layers of the superior colliculus were found to project strongly to the contralateral medial accessory olive immediately beside subnucleus and weakly to the same area ipsilaterally.Four pathways were identified as contributing fibers to the olivary projections. These were the medial longitudinal fasciculus, the medial tegmental tract, the central tegmental tract and tectospinal or tectobulbar fibers. The rubrospinal tract did not contribute projections to the olive. Injections in the caudate nucleus, entopeduncular nucleus and ventral anterior and ventral lateral thalamic nuclei, did not result in any labeling in the olive.List of Abbreviations AC anterior commissure - Cd caudate nucleus - CG central gray - CP cerebral peduncle - CTT central tegmental tract - DAO dorsal accessory olive - dc dorsal cap of Kooy - dmcc dorsomedial cell column of the inferior olive - dlPO dorsal lamella of the principal olive - Entop entopeduncular nucleus - EW nucleus of Edinger-Westphal - FR fasciculus retroflexus - Fx fornix - GP globus pallidus - H H field of Forel - HRP horseradish peroxidase - IC inferior colliculus - INC interstitial nucleus of Cajal - Int Cap internal capsule - IPN interpeduncular nucleus - LRN lateral reticular nucleus - MAO medial accessory olive - MB mammillary body - MGB medial geniculate body - MLF medial longitudinal fasciculus - MRF mesencephalic reticular formation - MTT medial tegmental tract - ND nucleus of Darkschewitsch - NFF nucleus of the fields of Forel - NPC nucleus of posterior commissure - NPP posterior pretectal nucleus - NRTP nucleus reticularis tegmenti pontis - n III third cranial nerve fibers - OT optic tract - PC posterior commissure - PF parafascicular nucleus - PG pontine gray - PO principal olive - PTM medial pretectal nucleus - RNp parvocellular red nucleus - RN red nucleus - RST rubrospinal tract - subnucleus beta of the inferior olive - sPf subparafascicular nucleus - SC superior colliculus - TH thalamus - vlPO ventral lamella of the principal olive - vlo ventral lateral outgrowth of the principal olive - VTA ventral tegmental area of Tsai - ZI zona incerta - III nucleus of third cranial nerve - XII nucleus of twelfth cranial nerve Supported by a grant from the Canadian Medical Research Council to the Group in Neurological Sciences at the Université de MontréalSupported by a postdoctoral fellowship of the Centre de Recherche en Sciences Neurologiques of the Université de Montréal     

Brain stem projections to the pulvinar-lateralis posterior complex of the cat

Summary The present experiments were undertaken to define the areas of projection of pretectum and superior colliculus to the pulvinar and n. lateralis posterior, respectively, and to define other brain stem structures projecting to these thalamic nuclei in cats. For this purpose the technique of retrograde transport of horseradish peroxidase (HRP) has been used.After injection of the enzyme in the pulvinar, neurons were labeled in all subdivisions of the pretectal area. The majority of the labeled cells were located in the n. pretectalis posterior and n. tractus opticus although cells filled with HRP were present also in the n. pretectalis anterior pars compacta and area pretectalis medialis. Neurons projecting to the pulvinar were also found in the periaqueductal gray, reticular formation and locus coeruleus.When HRP was injected in the n. lateralis posterior, labeled neurons were present in the II and III subdivisions of the second layer of the superior colliculus. The location of these cells shifted from medial to lateral as the injections were shifted from posterior to anterior within the lateralis posterior. Neurons projecting to this nucleus were also present in the intermediate layers of the superior colliculus, lateral hypothalamus and parabigeminal nucleus.The possible role of the pretectal area and superior colliculus in mediating somesthetic input to the pulvinar and lateralis posterior, respectively, and the role of these structures in the control of ocular movements, are discussed.Abbreviations APM area pretectalis medialis - Cu nucleus cuneiformis - CS nucleus centralis superior - fr fasciculus retroflexus - Gp pontine gray - Hb nucleus habenulae - IC inferior colliculus - LC locus coeruleus - LGB lateral geniculate body - LP lateralis posterior - MGB medial geniculate body - nPA_c nucleus pretectalis anterior pars compacta - nPA_r nucleus pretectalis anterior pars reticularis - nPC nucleus posterior commissurae - nPP nucleus pretectalis posterior - nTO nucleus tractus opticus - PAG periaqueductal gray - PB nucleus parabigeminalis - Pi pulvinar inferior - PO nucleus posterior of the thalamus - Pul pulvinar - Pt pretectum - RF reticular formation - Rtp tegmental reticular nucleus - SC superior colliculus Supported by H. de Jur Foundation and USPHS Grant TWO 2718Present address: Max-Planck-Institut für biophysikalische Chemie, Postfach 968, D-3400 Göttingen, Federal Republic of Germany     

Bulbar neurones with axonal projections to the trigeminal motor nucleus in the cat

Summary The location of bulbar neurones with axons projecting to the ipsi- and contralateral trigeminal motor nucleus were investigated in cats anaesthetized with sodium pentobarbital. Wheat germ agglutinin-conjugated horseradish peroxidase (WGA-HRP) was injected in amounts of 5–24 nl. A volume-calibrated microelectrode was used for recording of evoked potentials and pressure injection of WGA-HRP. The injection site was guided by the position where a maximal antidromic response was evoked by electrical stimulation of the masseteric nerve. The survival time was 19–22 h. In preparations with the depot located in the masseteric subnucleus retrogradely stained neurones were found bilaterally in the borderzone of the trigeminal motor nucleus. Dense populations of stained neurones were observed ipsi- and contralaterally in the dorsal division of the main sensory trigeminal nucleus and the subnucleus- of the oral nucleus of the spinal trigeminal tract. Clusters of WGA-HRP-neurones were observed bilaterally in the lateral tegmental field at the level of the subnucleus- of the oral nucleus of the spinal trigeminal tract, bilaterally dorsal to the facial nucleus and contralaterally adjacent to the hypoglossal nucleus. No stained neurones were found in the gigantocellular reticular nucleus. A group of stained neurones was located in the marginal nucleus of brachium conjunctivum and some were found in the raphé nuclei near obex. Cell profiles were of two types: medium-sized neurones with a triangular profile and 30–40 m diameter, and fusiform neurones 10×50–70 m. Convergence of descending cortical and trigeminal afferent inputs on interneurones located in the lateral borderzone of the trigeminal motor nucleus, i.e. the intertrigeminal area, is reported in the preceding paper.List of Abbreviations BCM Marginal nucleus of the brachium conjunctivum - CAE Nucleus caeruleus - CI Inferior central nucleus - Cu Cuneate nucleus - Cux External cuneate nucleus - DMV Dorsal motor nucleus of the vagus - FTG Gigantocellular tegmental field - FTL Lateral tegmental field - FTP Paralemniscal tegmental field - Gr Gracile nucleus - Mb Medial borderzone of NVmt - NintV Intertrigeminal area - NsV Supratrigeminal nucleus (area) - NVmes Mesencephalic trigeminal nucleus - NVmt Trigeminal motor nucleus - NVsnpr Main sensory trigeminal nucleus - NVsnpr-d Main sensory trigeminal nucleus, dorsal division - NVsnpr-v Main sensory trigeminal nucleus, ventral division - NVspc Caudal nucleus of the spinal trigeminal tract - NVspo- Subnucleus- of the oral nucleus of the spinal trigeminal tract - NVspo- Subnucleus- of the oral nucleus of the spinal trigeminal tract - NVspo- Subnucleus- of the oral nucleus of the spinal trigeminal tract - V Spinal trigeminal tract - NVII Facial nucleus - VII Facial nerve - NXII Hypoglossal nucleus - XII Hypoglossal nerve - Ols Superior olive - Rb Rostral borderzone of NVmt - Vb Ventral borderzone of NVmt - VIN Inferior vestibular nucleus - VSL Superior vestibular nucleus, lateral division     

Input-output organization of reticulospinal neurones,with special reference to connexions with dorsal neck motoneurones in the cat

Summary Dorsal neck motoneurones receive disynaptic tectal and pyramidal EPSPs via common reticulospinal neurones (RSNs). This study was aimed at identification of the RSNs projecting directly to neck motoneurones and mediating these EPSPs. 1. Stimulation of the tectum and the cerebral peduncle evoked monosynaptic descending volleys in the spinal cord, which were chiefly mediated by reticulospinal neurones in the pons and the medulla. Systematic tracking of the C3 and C7 segments was made to locate descending volleys in the spinal funiculi. The tectal monosynaptic volley was largest in the medial part of the ventral funiculus and decreased gradually as the recording electrode was moved to the lateral part of the ventral funiculus and the lateral funiculus. In contrast, the peduncle-evoked monosynaptic volley was distributed rather evenly in the ventral funiculus and the ventral half of the lateral funiculus. 2. Differences in funicular distribution of the two descending volleys suggest the existence of subgroups of RSNs which differed in strength of inputs from the two descending fibre systems and in the funicular location of descending axons. 3. The RSNs were classified into the following four groups; (1) mRSNs which descended in the medial part of the ventral funiculus, (2) in RSNs which descended in the ventrolateral funiculus, (3) 1RSNs which descended in the dorsal 2/3 of the lateral funiculus and (4) coRSNs which descended in the contralateral funiculi. The mRSNs were located in a fairly localized region corresponding to the nucleus reticularis pontis caudalis (N.r.p.c.), while inRSNs, 1RSNs and coRSNs were mainly in the nucleus reticularis gigantocellularis (N.r.g.), in the nucleus reticularis magnocellularis (N.r.m.) and in the nucleus reticularis ventralis (N.r.v.). RSNs were further divided into three types depending on the levels of projection. L-RSNs projected to the lumbar spinal segments. C-RSNs descended to the C6–C7 spinal segment but not to the lumbar segments. N-RSNs projected to the C3 but not to the C6–C7 segments. 4. Stimulation of the tectum and the cerebral peduncle produced monosynaptic negative field potentials in the medial two thirds of the reticular formation in the pons and medulla. Tectal field potentials were largest in the N.r.p.c. and the rostral part of the N.r.g., while pyramidal field potentials were largest in the N.r.g. Correspondingly, RSNs in the N.r.p.c. (mRSNs) received larger monosynaptic EPSPs from tectal than from pyramidal volleys, while RSNs in the N.r.g. (in-, 1- and coRSNs) received stronger input from the peduncle than from the tectum. 5. Stimulation of the C7 ventral but not the lateral funiculus evoked monosynaptic EPSPs on all the dorsal neck motoneurones tested. Stimulation of the L1 segment only produced monosynaptic EPSPs in 35% of the motoneurones. The L1 evoked EPSPs were much smaller than C7 evoked EPSPs. 6. The C7 evoked EPSPs (C7 EPSP) showed complete occlusion (collision) with the tectal or pyramidal disynaptic EPSPs. Similar results were obtained with L1 EPSPs. These results indicate that tectal and pyramidal disynaptic EPSPs in dorsal neck motoneurones were mediated chiefly by C-mRSNs and C-inRSNs and partly by L-RSNs.     

Descending projections from brainstem and sensorimotor cortex to spinal enlargements in the cat

Summary Single and double retrograde tracer techniques were employed in cats to investigate: (1) the topographical relationships between supraspinal neurons projecting to either the brachial or lumbosacral enlargement, (2) the distribution and relative frequency of single supraspinal neurons which project to both enlargements by means of axonal branching.In one group of cats large injections of horseradish peroxidase (HRP) were made throughout either the brachial or lumbosacral enlargement. The results from these experiments support recent observations on the multiplicity of brainstem centers giving origin to descending spinal pathways and provide evidence for a population of corticospinal neurons in area 6.In a second set of experiments, HRP was injected in one enlargement, and ³H-apo-HRP (enzymatically inactive) was injected in the other enlargement. Relatively large numbers of neurons with collateral projections to both enlargements (double-labeled) were observed in the medullary and pontine reticular formation, the medial and inferior vestibular nuclei bilaterally, the ipsilateral lateral vestibular nucleus, Edinger-Westphal nucleus, caudal midline raphe nuclei and nuclear regions surrounding the brachium conjunctivum. By contrast, double-labeled neurons were infrequently observed in the red nucleus and sensorimotor cortex, contralateral to the injections.In the red nucleus, lateral vestibular nucleus and sensorimotor cortex, neurons projecting to the brachial enlargement were largely segregated topographically from neurons projecting to the lumbosacral enlargement. However, there was some overlap, and double-labeled neurons were consistently observed within the region of overlap. In the sensorimotor cortex, the overlap between brachial- and lumbar-projecting neurons was most prominent in areas 4 and 3a, along the cruciate sulcus, but also involved other cytoarchitectonic regions in the medial aspect of the hemisphere.Abbreviations AM nucleus ambiguus - ap area postrema - aq aqueduct - BC brachium conjunctivum - ci central inferior nucleus of the raphe - cs central superior nucleus of the raphe - Cun cuneate nucleus - EC external cuneate nucleus - EW Edinger-Westphal nucleus - ETC central tegmental field - FTG gigantocellular tegmental field - FTL lateral tegmental field - FTM magnocellular tegmental field - FTP paralemniscal tegmental field - Gr gracile nucleus - IO inferior olive - K-F Kölliker-Fuse nucleus - LC nucleus locus coeruleus - li rostral linear nucleus of the raphe - LR lateral reticular nucleus - mlf medial longitudinal fasciculus - PAG periaqueductal gray - PbL lateral parabrachial nucleus - PG pontine gray - PON preolivary nucleus - ppr post-pyramidal nucleus of the raphe - RB restiform body - RNm red nucleus, magnocellular division - RNp red nucleus, parvocellular division - SC superior colliculus - SN substantia nigra - SOl lateral nucleus of the superior olive - SOm medial nucleus of the superior olive - Spin V spinal trigeminal nucleus - SubC nucleus subcoeruleus - TB trapezoid body - tb nucleus of the trapezoid body - trm tegmental reticular nucleus - VInf inferior vestibular nucleus - VLd lateral vestibular nucleus, dorsal division - VLv lateral vestibular nucleus, ventral division - VM medial vestibular nucleus - VSm superior vestibular nucleus, medial division Cranial Nerves and their Nuclei III oculomotor nucleus or nerve - V sensory nucleus of the trigeminal nerve - VI abducens nucleus - VII I facial nucleus, lateral part - VII m facial nucleus, medial part - X vagus nucleus - XII hypoglossal nucleus The research was supported by USPHS grants NS 12440 and MH 14277. ³H-apo-HRP was generously provided by New England Nuclear     

Widespread cortical projections of the ventral tegmental area and of other brain stem structures in the cat

Summary Thirty-three cat brains with injections of horseradish peroxidase in various regions of the cerebral cortex were screened for afferent projections from the ventral tegmental area, the locus ceruleus, and the parabrachial nuclei. All three structures were found to project to rather divergent parts of the cortex, including regions in the posterior half of the hemisphere. These results, especially for the ventral tegmental area and, to a lesser degree, for the parabrachial neurons, disagree with most of the target loci of established cortical afferents in the rat. Though our results might be attributed to species differences in the cortical innervation of brain stem structures, we prefer explanations which emphasize different densities in the distribution of brain stem afferents to the cortex, and/or which suggest different cortical targets of catecholaminergic and noncatecholaminergic neurons.Supported in part by grant Ma 795 from the Deutsche Forschungsgemeinschaft (DFG)     

An electrophysiological investigation of propriospinal inspiratory neurons in the upper cervical cord of the cat

Summary This study was performed in order to describe the location, axonal projection and possible synaptic action of the inspiratory neurons recently described in the upper cervical cord. In 26 cats anaesthetized with Nembutal, extracellular recordings were made from 224 cervical inspiratory units which were found near the lateral border of lamina VII and formed a column extending from the caudal end of the nucleus retroambigualis at the C₁ segment to the rostral half of the C₃ segment. Most of the units (approximately 85%) could be excited antidromically from the thoracic cord. Antidromic mapping showed collateral branches to the C₅ segment in the vicinity of the phrenic nucleus, occasionally crossing the midline. No synaptic connections with phrenic motoneurones could be revealed either by cross-correlation of the activity of the cervical units with the discharge of C₅ phrenic root, or by spike-triggered averaging (STA) of the post-synaptic noise recorded intracellularly from phrenic motoneurons. Extensive branching was found in the examined T₃–T₅ segments with arborizations near the ipsilateral intercostal motor nuclei and often extending across the midline. Cross-correlation experiments did not show clear monosynaptic connections to the inspiratory intercostal motoneurons. Intracellular recording from intercostal motoneurons and STA resulted in a few (2 out of 37) small, probably disynaptic, e.p.s.p.s. It is concluded that the upper cervical neurons are involved in the control of phrenic and intercostal motoneurons, probably through a disynaptic pathway involving segmental interneurons.