Neuronal mechanisms of the interaction of Deiters' nucleus with some brainstem structures

The effects of stimulation of the interstitial nucleus of Ramón y Cajal, as well as the nucleus of Darkschewitsch, inferior olive and nucleus reticularis tegmenti pontis on the neuronal activity of the lateral vestibular nucleus of Deiters were studied by means of an intracellular recording technique. Stimulation of these structures is shown to lead to antidromic and orthodromic activation of Deiters' neurons. Collaterals of vestibulospinal neurons entering these structures are revealed electrophysiologically. It was shown that stimulation of the rostral part of the inferior olive evoked in Deiters' neurons predominantly antidromic responses, whereas stimulation of the caudal part of the inferior olive leads mostly to synaptic activation. Stimulation of Ramón y Cajal's and Darkschewitsch's nuclei evokes mono- and/or polysynaptic excitatory and inhibitory postsynaptic potentials in Deiters' neurons. Mono-, oligo- and/or polysynaptic inhibitory postsynaptic potentials were also evoked by stimulation of nucleus reticularis tegmenti pontis, as well as the rostral and particularly, caudal parts of the inferior olive. Stimulation of the caudal part of the inferior olive evoked mono-, oligo- and/or polysynaptic excitatory postsynaptic potentials in Deiters' neurons. Convergence of influences from the stimulated structures on the vestibular neurons under study was shown. A topical correlation between Deiters' nucleus and the brainstem nuclei mentioned above was found. The presence of inhibitory and excitatory interaction of these structures with Deiters' nucleus was established.     

The lateral vestibular nucleus of the toadfish Opsanus tau: Ultrastructural and electrophysiological observations with special reference to electrotonic transmission

The lateral vestibular nucleus of the toadfish Opsanus tau was localized by means of axonal iontophoresis of Procion Yellow. The ultrastructure of the lateral vestibular nucleus neurons was then correlated with their electrophysiological properties. The lateral vestibular nucleus consists of neurons of various sizes which are distributed in small clusters over a heavily myelinated neuropil. The perikarya and main dendrites of the large and the small neurons are surrounded by a synaptic bed, which is separated from the neighboring neuropil by a layer of thin astrocytic processes. The synaptic bed contains three main classes of axon terminals, club endings, large and small terminals, the first being quite infrequent. All the large terminals as well as the occasionally observed club endings contain a pure population of rounded synaptic vesicles. In some of the small axon terminals there are also rounded vesicles; however, the majority contain flattened vesicles or a pleomorphic population. These data indicate that the small terminals originate from different afferent sources. The synaptic interfaces of the large boutons and of the club endings bear three types of junctional complexes: attachment plates, gap junctions and active zones. Those showing both gap junctions and active zones were designated as morphologically ‘mixed synapses’. Gap junctions, although in large number, have only been observed at the synaptic interfaces between terminals with rounded vesicles and the perikarya or the dendrite of the lateral vestibular nucleus neurons. Therefore electrotonic coupling would only be possible by way of presynaptic fibers. Some axons observed in the neuropil were found to establish gap junctional complexes with two different dendritec profiles and this observation is in favour of electrotonic coupling by way of presynaptic terminals.Field and intracellular potentials were recorded in the lateral vestibular nucleus. The field potential evoked by stimulation of the vestibular nerve consisted of an early positive-negative wave followed by a slow negativity, and that evoked by spinal cord stimulation was composed of an antidromic potential followed by a slow negative wave. Vestibulo-spinal neurons were identified by their antidromic spikes. In these cells, stimulation of the ipsilateral vestibular nerve evoked an excitatory postsynaptic potential with two components. The short delay of the first component of this excitatory postsynaptic potential and its ability to follow paired stimulation at close intervals without reduction of the second response suggest that it is transmitted electrotonically from primary vestibular afferent fibers. By contrast the latency of the second peak of the vestibular evoked excitatory postsynaptic potential and its sensitivity to high stimulus frequencies are compatible with monosynaptic chemically mediated transmission from primary vestibular afferents. Spinal stimulation evoked graded antidromic depolarizations in vestibulo-spinal neurons. The latency of these potentials was too short to allow for chemical transmission through afferents or recurrent collaterals and suggests electrotonic spread of antidromic activity from neighboring neurons. An important finding is that the graded antidromic depolarizations can initiate spikes; thus coupling between neurons in the lateral vestibular nucleus is sufficiently close that a cell can be excited by activity spread from neighboring cells. Similar graded depolarizations were recorded in identified primary vestibular afferents; their latencies and time course indicate that they were brought about by electrotonic spread of postsynaptic potentials and spikes to the impaled presynaptic fibers; this confirms the morphological evidence that coupling between lateral vestibular nucleus neurons occurs, at least in part, by way of presynaptic vestibular axons. As the spinal stimulus strength was increased, these graded depolarizations became large enough to initiate spikes which presumably propagate to the vestibular receptors. Thus antidromic invasion of the presynaptic terminals may provide negative feedback by preventing their re-excitation at short intervals after a synchronous discharge of an adequate number of postsynaptic cells. Excitatory inputs to the neurons of the lateral vestibular nucleus were identified from the spinal cord and from the contralateral vestibular nerve. Long latency excitatory postsynaptic potentials large enough to excite the cells were recorded following spinal stimulation; the threshold intensity for evoking them was consistently higher than that adequate to generate the graded antidromic depolarizations. Field potentials recorded after stimulation of the contra lateral vestibular nerve consisted of an initial positive negative wave followed by a slow negative wave. the stimulus intensity for evoking these potentials was the same or slightly above the threshold for those evoked in the lateral vestibular nucleus on the stimulated side. Also lateral vestibular nucleus neurons exhibited excitatory postsynaptic potentials large enough to excite the cells following stimulation of the contralateral vestibular nerve. but no inhibitory postsynaptic potentials were detected. This lack of commissural inhibition indicates a qualitative difference between the central organization of these cells in the toadfish and in mammals.The presence of neurons in the lateral vestibular nucleus which send their axons to the labyrinth was confirmed by their heavy staining with Procion Yellow following axonal iontophoresis. In a number of vestibular neurons. abruptly rising spikes were evoked at short latencies after adequate stimulation of the ipsilateral vestibular nerve. Graded stimuli applied to the vestibular nerve evoked graded short latency depolarizations as well as long latency excitatory postsynaptic potentials in these presumed efferent neurons to the labyrinth; the former could indicate electrotonic coupling of the efferent cells or electrotonic transmission from primary afferents, resulting in a short latency feedback loop.From these studies, the synaptic organization of the lateral vestibular nucleus neurons is compared with that of the Mauthner cells of teleosts, and the possibility of a dual mode of transmission, electrical and chemical, by primary vestibular afferents is discussed.     

Direct projections from vestibular nuclei to facial nucleus in cats

Postsynaptic potentials were recorded from motoneurons in the facial nucleus in response to stimulation of the vestibular and trigeminal nerves. The motoneurons were identified by antidromic activation from their peripheral axons. Disynaptic excitatory and inhibitory postsynaptic potentials (EPSPs and IPSPs) and mixed EPSP/IPSPs were recorded in response to vestibular nerve stimulation, ranging in latency from 0.9 to 2.1 ms, with most at 1.5 ms. Activity in secondary vestibular axons recorded within the facial nucleus occurred at a latency of 0.7-1.1 ms. The amplitudes of the vestibular postsynaptic potentials were small, generally less than a millivolt, but double shocks produced marked summation. The average time to peak of ipsilateral vestibular EPSPs, 1.1 ms, was faster than that of either ipsilateral IPSPs, 1.6 ms, or contralateral EPSPs, 1.4 ms. The double-spiked vestibular activity was detectable in double-peaked PSPs. Disynaptic EPSPs, ranging in latency from 2.0 to 3.0 ms, were recorded in response to trigeminal nerve stimulation. The average time to peak was 1.3 ms. The multiple-spiked activity of the trigeminal neurons was detectable in multipeaked EPSPs. Inhibitory ipsilateral effects (Vi IPSPs) were recorded twice as often as excitatory ipsilateral effects (Vi EPSPs), being found in 29% versus 15% of the motoneurons. Contralateral effects were found in 13% of the motoneurons studied, and almost all were excitatory. Analysis of synaptic potential shapes suggested that the excitatory and inhibitory vestibular synapses probably contact distal dendrites preferentially, with the excitatory connections being somewhat closer to the soma. The trigeminal inputs probably contact the facial motoneurons more extensively near the soma. Horseradish peroxidase was injected into the facial nucleus, and retrograde uptake by vestibular neurons was studied. The majority of filled vestibular neurons was ipsilateral to the injection site, especially in the medial vestibular nucleus, ventral y group, and supravestibular nucleus. On the contralateral side, filled vestibular cells were found almost exclusively in the medial nucleus. Filled cells were also noted in the trigeminal nucleus, predominantly ipsilaterally at all rostrocaudal levels. We have demonstrated monosynaptic projections to facial motoneurons from both vestibular and trigeminal nuclei. The trigeminal input is likely to be involved in facial reflexes, especially blinking and grimacing. The afferent vestibular population overlaps that going to the oculomotor and cervical motoneurons; these projections may be collaterals of single vestibular neurons.4+.     

Direct excitatory and inhibitory synaptic inputs from the medial mesodiencephalic junction to motoneurons innervating extraocular oblique muscles in the cat

Summary This study investigated the nature of synaptic inputs from the Forel's field H (FFH) in the medial mesodiencephalic junction to inferior oblique (IO) motoneurons in the oculomotor nucleus and superior oblique (SO) motoneurons in the trochlear nucleus in anesthetized cats, using intracellular recording techniques. Stimulation of the FFH induced monosynaptic EPSPs in IO motoneurons on both sides. Paired stimulation of the ipsilateral FFH and contralateral vestibular nerve substantiated that the FFH-induced EPSPs were caused mainly by direct excitatory fibers from the FFH to IO motoneurons and partly by axon collaterals of excitatory neurons in the vestibular nuclei. Among parts of the FFH, the medial part was most effective for producing the EPSPs. Systematic tracking with the stimulating electrode in and around the FFH revealed that effective sites of stimulation inducing negative field potentials in the IO subdivision of the oculomotor nucleus, identified as extracellular counterparts of the EPSPs in IO motoneurons, were also located in the interstitial nucleus of Cajal, nearby reticular formation and posterior commissure, besides within and near the medial part of the FFH. Areas far rostral, dorsal and ventral to the FFH were ineffective. EPSP-IPSPs or EPSPs were mainly induced in SO motoneurons on both sides by FFH stimulation. Latencies of these EPSPs and IPSPs were close to those of the EPSPs in IO motoneurons, indicating their monosynaptic nature. Effective stimulation sites for inducing these synaptic potentials overlapped those for the EPSPs in IO motoneurons. Based on these results, it was suggested that excitatory and inhibitory premotor neurons directly controlling IO and SO motoneurons were located within and near the medial part of the FFH.     

The neuronal organization of horizontal semicircular canalactivated inhibitory vestibulocollic neurons in the cat

Summary 1.The somatic location and axonal projections of inhibitory vestibular nucleus neurons activated by the horizontal semicircular canal nerve (HCN) were studied in anesthetized cats. Cats were anesthetized with ketamine hydrochloride and pentobarbital sodium. 2.Intracellular recordings were obtained from 11 neck extensor motoneurons which were identified by antidromic activation from the dosai rami (DR) in the C₁ segment. Stimulation of the ipsilateral (i-) HCN and the ipsilateral abducens (AB) nucleus evoked IPSPs in the motoneurons. These IPSPs were fully or partially occluded when they were evoked simultaneously. 3. Intracellular recordings were obtained from 8 AB motoneurons. Stimulation of the i-HCN and the i-C₁DR motoneuron pool evoked IPSPs in the AB motoneurons. These IPSPs were also partially occluded when they were evoked simultaneously, which implied that some HCN-activated neurons inhibit both i-AB motoneurons and ipsilateral neck motoneurons. 4. Unit activity was extracellularly recorded from 30 vestibular neurons that were activated monosynaptically by i-HCN stimulation. Their axonal projections were determined by stimulating the i-AB nucleus and the i-C₁DR motoneuron pool. Eight neurons were activated by both stimuli, and were termed vestibulooculo-collic (VOC) neurons. Their axonal branching was examined by means of local stimulation in and around the i-AB nucleus and the i-C₁DR motoneuron pool. Eighteen neurons were antidromically activated from the i-C₁DR motoneuron pool but not from the i-AB nucleus. These were termed vestibulo-collic (VC) neurons. Four neurons were activated from the i-AB nucleus but not from the ventral funiculus in the C₁ segment, and were termed vestibulo-ocular (VO) neurons. The HCN-activated inhibitory neurons were mostly localized in the rostroventral part of the medial vestibular nucleus. 5. Horseradish peroxidase (HRP) was injected iontophoretically into descending axons of 2 HCN-activated inhibitory VOC neurons which were identified by stimulation of the i-HCN and the i-AB nucleus. Axon collaterals were ramified from a stem axon in the ventral funiculus, and entered the gray matter and spread in the laminae VIII and IX. Terminal boutons were distributed over the medial and the ventromedial parts of the vental horn in the C₁ segment.     

Synaptic mechanisms of interaction between Deiters' nucleus and the nuclei of some cranial nerves

The effects of stimulation of the vestibular nerve, spinal trigeminal nucleus, facial and hypoglossal nuclei of the cranial nerves on the neuronal activity in the lateral vestibular nucleus of Deiters were studied in cats anaesthetized with pentobarbitone. Stimulation of these nuclei was found to produce antidromic and synaptic activation of Deiters' neurons. Descending axon collaterals of the vestibular neurons to these brainstem structures were revealed. Stimulation of the VIIIth nerve, spinal trigeminal and facial nuclei evoked mono- and polysynaptic excitatory postsynaptic potentials in Deiters' neurons. Stimulation of the spinal trigeminal nucleus evoked mono- and polysynaptic inhibitory postsynaptic potentials and disfacilitation in Deiters' neurons. In some vestibular neurons inhibitory postsynaptic potentials were also evoked by stimulation of the nucleus hypoglossus. Convergence of influences from these structures on Deiters' neurons was shown to exist. The peculiarities and functional significance of the effects mentioned are discussed.     

Properties of ventromedial hypothalamic neurons with axons to midbrain central gray

Summary In female rats anesthetized with urethane, 151 neurons in and around the ventromedial nucleus of the hypothalamus were identified by antidromic activation as having axonal projection to the mesencephalic central gray at the midcollicular level. Identified neurons were most numerous in the rostral part and at the borders of the nucleus.Antidromic spike latencies, constant for a given cell to stimulation with fixed intensity at a low repetition rate, had a wide range across cells (1.4–41.5 ms). In 37 cells, gradual increases in stimulus intensity allowed sudden discrete latency decreases as large as 9.8 ms. These may reflect activation of separate axonal branches of terminal arborizations.Eleven among 43 tested cells were antidromically driven from the dorsal longitudinal fasciculus (DLF) at the diencephalic-mesencephalic junction as well as from the central gray. Latencies of DLF responses were always shorter than those from central gray. From this and collision experiments between central gray-evoked and DLF-evoked antidromic spikes, it was concluded that at least one quarter of mesencephalic projections from the ventromedial nucleus descend through DLF. The mean conduction velocity of these axons was 0.8 m/s, indicating that they belong to thin unmyelinated C-group fibers.Thirty percent of the cell population studied received excitatory input from the cortical or medial nucleus of the amygdala. Four cells were identified as having projections both to the central gray and the amygdala.Estrogen treatment of ovariectomized female rats caused no major changes in antidromic latency, absolute refractory period or resting activity of these identified hypothalamic neurons. However, the stimulation threshold for antidromic activation was significantly lower in the estrogen-treated animals.Axons to the central gray from ventromedial hypothalamic neurons provide for hypothalamic bias on brain stem reflex paths, for reproductive and other behaviors.Supported by NIH grant HD-05751 and by institutional grant from the Rockefeller Foundation for the study of reproductive biology     

Functional organization of premotor neurons in the cat medial vestibular nucleus related to slow and fast phases of nystagmus

Summary Extracellular spikes were recorded from secondary vestibular neurons in the cat medial vestibular nucleus (MVN) and were identified as type I or II neurons by horizontal rotation. Type I neurons were further classified as excitatory or inhibitory premotor neurons on the basis of their axonal termination in the contralateral or ipsilateral abducens nucleus, demonstrated by spike-triggered averaging of abducens nerve discharges, or by antidromic activation using systematic microstimulation within the abducens nucleus.Both excitatory and inhibitory premotor type I MVN neurons exhibited a rhythmic modulation of their firing rate in association with nystagmus elicited by rotation or electrical stimulation of the vestibular nerve. Their tonic activity during the slow phase was suppressed at the quick phase directed to the ipsilateral side.Excitatory type I MVN neurons terminating in the contralateral abducens nucleus sent collateral axons to the contralateral MVN. These commissural neurons also showed a nystagmus-related discharge pattern.Type II MVN neurons activated at short latency by stimulation of the contralateral vestibular nerve exhibited burst discharges when the activity of ipsilateral type I neurons was suppressed at the quick phase. These type II neurons made monosynaptic inhibitory connections with type I neurons as shown by the post-spike average of the membrane potential of secondary MVN neurons triggered from spikes of single type II neurons. Thus, the inhibitory action originating from burst activity of type II MVN neurons contributes to suppression of type I premotor MVN neurons during fast eye movements.Supported by Grant-in-Aid for Scientific Research No. 248106 from Japan Ministry of Education, Science and Culture. Dr. Schor was supported by the Research Fellowship of Japan Society for the Promotion of Science (JSPS)     

Saccular and utricular inputs to single vestibular neurons in cats

Saccular and utricular organs are essential for postural stability and gaze control. Although saccular and utricular inputs are known to terminate on vestibular neurons, few previous studies have precisely elucidated the origin of these inputs. We investigated the saccular and utricular inputs to single vestibular neurons in whole vestibular nuclei of decerebrated cats. Postsynaptic potentials were recorded from vestibular neurons after electrical stimulation of the saccular and utricular nerves. Ascending and descending axonal projections were examined by stimulating the oculomotor/trochlear nuclei and the cervical segment of the spinal cord, respectively. After each experiment, locations of recorded neurons were identified. The recorded neurons (140) were classified into vestibulo-spinal (79), vestibulo-oculo-spinal (9), and vestibulo-ocular (3) neurons based on antidromic responses; 49 other vestibular neurons were unidentified. The majority of recorded neurons were mainly located in the lateral vestibular nucleus. Most of the otolith-activated vestibular nuclei neurons seemed to participate in vestibulospinal reflexes. Of the total 140 neurons recorded, approximately one third (51) received saccular and utricular inputs (convergent neurons). The properties of these 51 convergent neurons were further investigated. Most (33/51) received excitatory postsynaptic potentials (EPSPs) after saccular and utricular nerve stimulation. These results implied that most of the convergent neurons in this study additively coded mixed information for vertical and horizontal linear acceleration. Based on the latencies of convergent neurons, we found that an early integration process for vertical and horizontal linear acceleration existed at the second-order level.     

Behavior of reticular,vestibular and prepositus neurons terminating in the abducens nucleus of the alert cat

Summary The activity of pontomedullary reticular, vestibular, and prepositus neurons has been recorded in the alert cat during spontaneous and vestibular-induced eye movements. Neurons were identified by their antidromic activation from the abducens nucleus. Spikes of these neurons were used to trigger the recording of field potentials in the abducens nucleus. The analysis by post-spike averaging of the field potentials showed the presence of a trifold system of reciprocal (excitatory and inhibitory) direct projections that originated in the above nuclei and terminated in the abducens nucleus with a distinctly graded effectiveness. This trifold afferent system is involved in the generation of fast eye movements, slow compensatory movements of vestibular origin, and eye fixation, respectively.     

Axonal trajectories of inhibitory vestibulocollic neurons activated by the anterior semicircular canal nerve and their synaptic effects on neck motoneurons in the cat

Summary Somatic location, axonal trajectories and synaptic effects of inhibitory vestibulocollic neurons which were activated by selective stimulation of the anterior semicircular canal nerve (ACN) were studied in the anesthetized cat. ACN stimulation evoked disynaptic inhibitory postsynaptic potentials (IPSPs) in neck flexor motoneurons. This was seen in all the (64/64) tested motoneurons innervating the ipsilateral (i-) longus capitis (LC) and the i-sternocleidomastoideus (SCM) muscles and in 86% (38/44) of the motoneurons innervating the contralateral (c-) LC muscle. The inhibitory relay neurons, identified by orthodromic and antidromic responses to stimulation of the ACN and the i- and c-LC motoneuron pools, were classified as VCi (vestibulocollic neurons sending an axon to the i-LC motoneuron pool) and VCc (vestibulocollic neurons sending an axon to the c-LC motoneuron pool) neurons. Neither VCi nor VCc neurons were activated antidromically by localized stimulation of the ascending medial longitudinal fasciculus (asc. MLF) or the 3rd nuclei. They were located in the medial, descending and ventral lateral vestibular nuclei. It was also observed that VCi neurons produced unitary IPSPs in i-LC and i-SCM motoneurons in the C1 segment. Inhibitory synapses were estimated to be on the cell somata and/or the proximal dendrites of the motoneurons.     

Electrophysiological properties of the somatotopic organization of the vestibulospinal system in the frog

In experiments on the preparation of a frog perfused brain (Rana ridibunda), field and intracellular potentials were recorded from neurons of the vestibular nuclear complex following stimulation of the ipsilateral vestibular nerve and different levels of the spinal cord. Stimulation of the vestibular nerve evoked mono- and polysynaptic excitatory postsynaptic potentials and orthodromic action potentials. In parallel, an antidromic activation of vestibular neurons sending their axons to the labyrinth was recorded. Vestibulospinal neurons sending their axons to the cervical (C neurons) and lumbar (L neurons) enlargements of the spinal cord were identified by their antidromic activation. A rather high conduction velocity along vestibulospinal fibres (mean 15.47 m/s) was observed. A somatotopic arrangement of the vestibulospinal system was established in spite of extremely large overlapping zones for the fore- and hindlimb representations in the vestibular nuclear complex. The hindlimbs were represented more poorly than the forelimbs. Antidromic potentials of C and L neurons were recorded in the medial, descending and with the highest density in the lateral vestibular nuclei (Deiters' nucleus). C neurons were evenly distributed in the other vestibular nuclei studied, while L neurons were located predominantly in the caudal parts of the vestibular nuclear complex. The multiplicity of the origin of the vestibulospinal axons was established. Peculiarities of the functional correlation between the vestibular input and vestibulospinal system are discussed.     

Axon collaterals of anterior semicircular canal-activated vestibular neurons and their coactivation of extraocular and neck motoneurons in the cat

We studied the ascending and descending axonal trajectories of excitatory vestibular neurons related to the anterior semicircular canal, by means of local stimulation and spike-triggered signal averaging techniques in anesthetized cats. More than 200 vestibular neurons related to the ampullary nerve of the anterior semicircular canal (ACN) were identified as vestibulo-ocular neurons by antidromic stimulation of the contralateral inferior oblique (IO) muscle motoneuron pool. In the descending, medial and ventral lateral nuclei, about 60% of these vestibulo-ocular neurons were also activated antidromically by upper cervical spinal cord stimulation (vestibulo-ocular-collic (cervical) = VOC). These VOC neurons produced unitary EPSPs in the majority of neck extensor motoneurons located at the C₁ segment. None of the VOC neurons had axons descending as far as the thoracic level. Most of these VOC neurons were activated monosynaptically following stimulation of the ACN. The conduction velocity of the descending axons of VOC neurons was approximately 63 m/s, which was significantly faster than that of the ascending axons. The remaining 40% of the vestibulo-ocular neurons were not activated antidromically following spinal cord stimulation at intensities of 1 mA or more (vestibulo-ocular = VO). Most of the VO neurons were activated polysynaptically by ACN stimulation. The superior vestibular nucleus contained VO neurons that were activated mono- and polysynaptically following ACN stimulation.     

Patterns of functional synaptic connections in organized cultures of cerebellum

Organized cultures of mouse cerebellum with separated regions containing cortical, deep nuclear neurons and brain stem neurons from the peduncular zone were used for electrophysiological studies of axonal projections and synaptic interactions. Responses of single neurons of each of the regions were recorded extracellularly and intracellularly during localized electrical stimulation of other parts of the explant, and indicated extensive synaptic interactions. Cortical stimulation inhibited deep nuclear neurons, apparently monosynaptically, and frequently caused antidromic activation of these cells. Synaptic responses of brain stem neurons to cortical stimulation were usually excitatory, but these were often succeeded by inhibitory potentials. Since brain stem cells were often antidromically activated, the excitatory synaptic responses may be mediated by collaterals of antidromically stimulated brain stem axons. Stimulation of the deep nuclear region could evoke inhibitory or excitatory potentials in cortical neurons, the most frequent response being an excitatory postsynaptic potential which was followed in about 2 ms by an inhibitory potential. Most excitatory and some inhibitory postsynaptic potentials followed high frequency stimulation with constant latencies.The results indicate that within these cultures there are rich synaptic interconnections, many of which follow patterns resembling those seen in the intact brain. The monosynaptic inhibitory projection from the cortex to the deep nuclei and collateral inhibition by Purkinje cell axons appear to be features of cerebellar function that are reproduced in this culture model. In addition, a projection from the deep nuclei to the cortex recently described in the intact cerebellum is also present in the cultures and gives postsynaptic potential responses typical of excitatory afferents to the cerebellar cortex. Such cultures appear useful as an experimental model for the study of synaptic mechanisms or the effects of drugs in the mammalian CNS.     

Properties of utricular nerve-activated vestibulospinal neurons in cats

The axonal pathway, conduction velocities, and locations of the cell bodies of utricular nerve-activated vestibulospinal neurons were studied in decerebrated or anesthetized cats using the collision test of orthodromic and antidromic spikes. For orthodromic stimulation, bipolar tungsten electrodes were placed on the utricular nerve and the other vestibular nerve branches were transected. Monopolar tungsten electrodes were positioned on both sides of the upper cervical segments (C2–4), caudal end of the cervical enlargement (C7-T1), and from the lower thoracic to the upper lumbar segments (T12-L3) and were used for antidromic stimulation of the spinal cord. Another monopolar electrode was also placed in the oculomotor nucleus to study whether utricular nerve-activated vestibulospinal neurons have ascending branches to the oculomotor nucleus. Of the 173 vestibular neurons orthodromically activated by the stimulation of the utricular nerve, 46 were second-order vestibulospinal neurons and 5 were third-order neurons. The majority of the utricular nerve-activated vestibulospinal neurons were located in the rostral part of the descending vestibular nucleus and the caudal part of the ventral lateral nucleus. Seventy-three percent of the utricular nerve-activated vestibulospinal neurons descended through the ipsilateral lateral vestibulospinal tract. Approximately 80% of these neurons reached the cervicothoracic junction, but a few reached the upper lumbar spinal cord. Twenty-seven percent of the utricular nerve-activated vestibulospinal neurons descended through the medial vestibulospinal tract or the contralateral vestibulospinal tracts. Those axons terminated mainly in the upper cervical segments. Almost none of the utricular nerve-activated vestibular neurons had ascending branches to the oculomotor nucleus.     

Convergence of the anterior semicircular canal and otolith afferents on cat single vestibular neurons

The convergence between the anterior semicircular canal (AC) and utricular (UT) inputs, as well as the convergence between the AC and saccular (SAC) inputs in single vestibular neurons of decerebrated cats were investigated. Postsynaptic potentials were recorded intracellularly after selective stimulation of each pair of vestibular nerves AC/UT or AC/SAC. Neurons were recorded from the central parts of the vestibular nuclei, where the otolith afferents mainly terminate. Of a total of 105 neurons that were activated after stimulation of the AC and UT nerves, 42 received convergent inputs. Thirty-eight of these neurons received excitatory inputs from both afferents. Convergent neurons were further classified into vestibulospinal (n=28) and vestibulooculospinal (n=6) neurons by antidromic activation from the border between the C1 and C2 spinal cord and the oculomotor or trochlear nucleus. Eight neurons that were not antidromically activated from either site were classified as vestibular neurons. Forty three percent of the convergent vestibulospinal neurons and most of the convergent vestibulooculospinal neurons projected to the spinal cord through the medial vestibulospinal tract. The remaining vestibulospinal and vestibulooculospinal neurons descended through the ipsilateral lateral vestibulospinal tract. Of a total of 118 neurons that were activated after stimulation of the AC and/or SAC nerves, 51 received convergent inputs (27 vestibulospinal, 4 vestibulooculospinal, 5 vestibuloocular and 15 vestibular neurons). Forty-two of the convergent neurons received excitatory inputs from both afferents. Thirty seven percent of the convergent vestibulospinal neurons and all of the convergent vestibulooculospinal neurons projected to the spinal cord through the medial vestibulospinal tract. The remaining vestibulospinal and vestibulooculospinal neurons descended through the ipsilateral lateral vestibulospinal tract. Electronic Publication     

Projections and firing properties of down eye-movement neurons in the interstitial nucleus of Cajal in the cat

To clarify the role of the interstitial nucleus of Cajal (INC) in the control of vertical eye movements, projections of burst-tonic and tonic neurons in and around the INC were studied. This paper describes neurons with downward ON directions. We examined, by antidromic activation, whether these down INC (d-INC) neurons contribute to two pathways: a commissural pathway to the contralateral (c-) INC and a descending pathway to the ipsilateral vestibular nucleus (i-VN). Stimulation of the two pathways showed that as many as 74% of neurons were activated antidromically from one of the pathways. Of 113 d-INC neurons tested, 44 were activated from the commissural pathway and 40 from the descending pathway. No neurons were activated from both pathways. We concluded that commissural and descending pathways from the INC originate from two separate groups of neurons. Tracking of antidromic microstimulation in the two nuclei revealed multiple low-threshold sites and varied latencies; this was interpreted as a sign of existence of axonal arborization. Neurons with commissural projections tended to be located more dorsally than those with descending projections. Neurons with descending projections had significantly greater eye-position sensitivity and smaller saccadic sensitivity than neurons with commissural projections. The two groups of INC neurons increased their firing rate in nose-up head rotations and responded best to the rotation in the plane of contralateral posterior/ipsilateral anterior canal pair. Neurons with commissural projections showed a larger phase lag of response to sinusoidal rotation (54.6 +/- 7.6 degrees ) than neurons with descending projections (45.0 +/- 5.5 degrees ). Most neurons with descending projections received disynaptic excitation from the contralateral vestibular nerve. Neurons with commissural projections rarely received such disynaptic input. We suggest that downward-position-vestibular (DPV) neurons in the VN and VN-projecting d-INC neurons form a loop, together with possible commissural loops linking the bilateral VNs and the bilateral INCs. By comparing the quantitative measures of d-INC neurons with those of DPV neurons, we further suggest that integration of head velocity signals proceeds from DPV neurons to d-INC neurons with descending projections and then to d-INC neurons with commissural projections, whereas saccadic velocity signals are processed in the reverse order.     

Intracellular study of frog's vestibular neurons in relation to the labyrinth and spinal cord

Summary Field and intracellular potentials were recorded in the vestibular nuclei of the frog following stimulation of the anterior branch of the ipsilateral vestibular nerve and the spinal cord. The field potential induced by stimulation of the vestibular nerve consisted of an early positive-negative wave followed by a slow negativity and that recorded during spinal cord stimulation was composed of an antidromic potential followed by a slow negative wave. These potentials were most prominent in the ventral region of the stato-acoustic complex. Mono- and polysynaptic EPSPs were recorded from vestibular neurons following vestibular nerve stimulation. Short latency depolarizations of small amplitude preceded the monosynaptic EPSPs in some neurons. Spike-like partial responses were commonly superimposed on the EPSPs. These all-or-none depolarizations probably originated in the dendrites. In a group of vestibular neurons stimulation of the vestibular nerve evoked full action potentials with latencies ranging from 0.2 to 1.1 msec. They are presumably caused by antidromic activation of neurons which send their axons to the labyrinth. The presence of efferent neurons in the vestibular nuclei was confirmed by their successful staining with Procion Yellow following axonal electrophoresis.After stimulation of the spinal cord, antidromic spike potentials and EPSPs were recorded in vestibular neurons. In addition, short-latency depolarizing potentials (EDPs) were evoked by spinal stimulation, with latencies similar to those of antidromic potentials. The EDPs are suggested to be induced by electrotonic transmission from the neighboring cell and likely to be active spike potentials produced at some distance away from the soma.     

Excitatory connections between neurons of the central cervical nucleus and vestibular neurons in the cat

 The central cervical nucleus (CCN) of the cat receives input from upper cervical muscle afferents, particularly primary spindle afferents. Its axons cross in the spinal cord, and while in the contralateral restiform body give off collaterals to the vestibular nuclei. In order to study the connections between CCN axons and vestibular neurons, we stimulated the area of the CCN in decerebrate cats while recording intra- or extracellularly from neurons in the contralateral vestibular nuclei. CCN stimulation evoked excitatory postsynaptic potentials (EPSPs) or extracellularly recorded firing in the lateral, medial and descending vestibular nuclei. The latency of EPSPs (mean 1.6 ms) was on average 0.4 ms longer than the latency of antidromic spikes evoked in the CCN by stimulation of the contralateral vestibular nuclei (mean 1.2 ms), demonstrating that the excitation was typically monosynaptic. The results provide further evidence that the CCN is an important excitatory relay between upper cervical muscle afferents and neurons in the contralateral vestibular nuclei. Received: 1 August 1996 / Accepted: 16 December 1996     

Extracellular recording of vestibulo-thalamic neurons projecting to the spinal cord in the cat

Forty vestibulo-thalamic (VT) neurons were recorded extracellularly in the vestibular nuclei of the anesthetized cat. More than half of the VT neurons responded monosynaptically to vestibular nerve stimulation; the others responded polysynaptically. The VT neurons were activated antidromically from one or two sites in the contralateral VPL, VPM, VL, VM, SG, and PO in the thalamus. Their axonal arborizations in the thalamus were likely restricted in narrow areas. About three quarters of the VT neurons were also activated antidromically from the ventral funiculus in the C1 segment. Axonal branchings were found in the contralateral C1 gray matter. The VT neurons were mainly localized in the descending vestibular nucleus.