Convergence patterns of the posterior semicircular canal and utricular inputs in single vestibular neurons in cats

The convergence of the posterior semicircular canal (PC) and utricular (UT) inputs in single vestibular nuclei neurons was studied intracellularly in decerebrate cats. A total of 160 vestibular neurons were orthodromically activated by selective stimulation of the PC and the UT nerve and classified according to whether or not they were antidromically activated from the spinal cord and oculomotor nuclei into vestibulospinal (VS), vestibulooculospinal (VOS), vestibuloocular (VO), and unidentified vestibular neurons. Fifty-three (33%) of 160 vestibular neurons received convergent inputs from both the PC and UT nerves. Seventy-nine (49%) vestibular neurons responded to PC inputs alone, and 28 (18%) neurons received inputs only from the UT nerve. Of 53 convergent neurons, 8 (15%) were monosynaptically excited from both nerves. Thirty-five (66%) received monosynaptic excitatory inputs from the PC nerve and polysynaptic excitatory or inhibitory inputs from the UT nerve, or vice versa. Approximately one-third of VS and VOS neurons received convergent inputs. A majority of the VS neurons descended to the spinal cord through the lateral vestibulospinal tract, while almost all the VOS neurons descended to the spinal cord through the medial vestibulospinal tract. The convergent neurons were found in all vestibular nuclei but more in the lateral nucleus and descending nucleus. The VS neurons were more numerous than VO neurons or VOS neurons.     

Convergence of posterior semicircular canal and saccular inputs in single vestibular nuclei neurons in cats

Convergence between posterior canal (PC) and saccular (SAC) inputs in single vestibular nuclei neurons was investigated in decerebrated cats. Postsynaptic potentials were recorded intracellularly after selective stimulation of the SAC and PC nerves. Stimulation of either the SAC or PC nerve orthodromically activated 143 vestibular nuclei neurons. Of these, 61 (43%) were antidromically activated by stimulation of the C1-C2 junction, 14 (10%) were antidromically activated by stimulation of the oculomotor or trochlear nucleus, and 14 (10%) were antidromically activated by stimulation of both the oculomotor or trochlear nucleus and the spinal cord. Fifty-four (38%) neurons were not activated by stimulation of either or both. We named these neurons vestibulospinal (VS), vestibulo-ocular (VO), vestibulooculo-spinal (VOS) and vestibular (V) neurons, respectively. Both PC and SAC inputs converged in 47 vestibular nuclei neurons (26 VS, 2 VO, 6 VOS and 13 V neurons). Of these, 19 received monosynaptic excitatory inputs from both nerves. This input pattern was frequently seen in VS neurons. Approximately half of the convergent VS neurons descended to the spinal cord through the lateral vestibulospinal tract. The remaining half and all the convergent VOS neurons descended to the spinal cord through the medial vestibulospinal tract. Most of the convergent neurons were located in the lateral nucleus or descending nucleus.     

Properties of utricular-activated vestibular neurons that project to the contralateral vestibular nuclei in the cat

The properties of utricular (UT)-activated vestibular neurons that send axons to the contralateral vestibular nuclei (commissural neurons) were investigated intracellularly or extracellularly in decerebrate cats. A total of 27 vestibular neurons were orthodromically activated by stimulation of UT nerves and antidromically activated by stimulation of the contralateral vestibular nuclei. All neurons tested were classified as vestibulospinal (VS), vestibulooculospinal (VOS), vestibuloocular (VO), and unidentified vestibular neurons (V) after antidromic stimulation of the spinal cord and oculomotor/trochlear nuclei. Most UT-activated commissural neurons (20/27) received monosynaptic inputs. Twelve of 27 commissural neurons were located in the medial vestibular nucleus, 5 were in the lateral vestibular nucleus, 10 were in the descending vestibular nucleus, and no commissural neurons were recorded in the superior vestibular nucleus. Seven of 27 neurons were commissural VS neurons, 9 of 27 were commissural VOS neurons, and 11 of 27 were commissural V neurons. No commissural VO neurons were found. All VOS neurons and 3 VS neurons issued descending axons via the medial vestibulospinal tract. We also studied convergent inputs from the posterior semicircular canal (PC) nerve onto UT-activated commissural neurons. Five of 27 UT-activated commissural neurons received converging inputs from the PC nerves. Electronic Publication     

Otolith and canal integration on single vestibular neurons in cats

In this review, based primarily on work from our laboratory, but related to previous studies, we summarize what is known about the convergence of vestibular afferent inputs onto single vestibular neurons activated by selective stimulation of individual vestibular nerve branches. Horizontal semicircular canal (HC), anterior semicircular canal (AC), posterior semicircular canal (PC), utricular (UT), and saccular (SAC) nerves were selectively stimulated in decerebrate cats. All recorded neurons were classified as either projection neurons, which consisted of vestibulospinal (VS), vestibulo-oculospinal (VOS), vestibulo-ocular (VO) neurons, or non-projection neurons, which we simply term vestibular (V) neurons. The first three types could be successfully activated antidromically from oculomotor/trochlear nuclei and/or spinal cord, and the last type could not be activated antidromically from either site. A total of 1228 neurons were activated by stimulation of various nerve pair combinations. Convergent neurons were located in the caudoventral part of the lateral, the rostral part of the descending, and the medial vestibular nuclei. Otolith-activated vestibular neurons in the superior vestibular nucleus were extremely rare. A high percentage of neurons received excitatory inputs from two nerve pairs, a small percentage received reciprocal convergent inputs and even fewer received inhibitory inputs from both nerves. More than 30% of vestibular neurons received convergent inputs from vertical semicircular canal/otolith nerve pairs. In contrast, only half as many received convergent inputs from HC/otolith-nerve pairs, implying that convergent input from vertical semicircular canal and otolith-nerve pairs may play a more important role than that played by inputs from horizontal semicircular canal and otolith-nerve pairs. Convergent VS neurons projected through the ipsilateral lateral vestibulospinal tract (i-LVST) and the medial vestibulospinal tract (MVST). Almost all the VOS neurons projected through the MVST. Convergent neurons projecting to the oculomotor/trochlear nuclei were much fewer in number than those projecting to the spinal cord. Some of the convergent neurons that receive both canal and otolith input may contribute to the short-latency pathway of the vestibulocollic reflex. The functional significance of these convergences is discussed.     

Properties and axonal trajectories of posterior semicircular canal nerve-activated vestibulospinal neurons

We studied the axonal projections of vestibulospinal neurons activated from the posterior semicircular canal. The axonal projection level, axonal pathway, and location of the vestibulospinal neurons originating from the PC were investigated in seven decerebrated cats. Selective electrical stimulation was applied to the PC nerve, and extracellular recordings in the vestibular nuclei were performed. The properties of the PC nerve-activated vestibulospinal neurons were then studied. To estimate the neural pathway in the spinal cord, floating electrodes were placed at the ipsilateral (i) and contralateral (c) lateral vestibulospinal tract (LVST) and medial vestibulospinal tract (MVST) at the C1/C2 junction. To elucidate the projection level, floating electrodes were placed at i-LVST and MVST at the C3, T1, and L3 segments in the spinal cord. Collision block test between orthodromic inputs from the PC nerve and antidromic inputs from the spinal cord verified the existence of the vestibulospinal neurons in the vestibular nuclei. Most (44/47) of the PC nerve-activated vestibulospinal neurons responded to orthodromic stimulation to the PC nerve with a short (<1.4 ms) latency, indicating that they were second-order vestibulospinal neurons. The rest (3/47) responded with a longer (≥1.4 ms) latency, indicating the existence of polysynaptic connections. In 36/47 PC nerve-activated vestibulospinal neurons, the axonal pathway was histologically verified to lie in the spinal cord. The axons of 17/36 vestibulospinal neurons projected to the i-LVST, whereas 14 neurons projected to the MVST, and 5 to the c-LVST. The spinal segment levels of projection of these neurons elucidated that the axons of most (15/17) of vestibulospinal neurons passing through the i-LVST reached the L3 segment level; none (0/14) of the neurons passing through the MVST extended to the L3 segment level; most (13/14) of them did not descend lower than the C3 segment level. In relation to the latency and the pathway, 33/36 PC nerve-activated vestibulospinal neurons were second-order neurons, whereas the remaining three were polysynaptic neurons. Of these, 33 second-order vestibulospinal neurons, 16 passed through the i-LVST, while 13 and 4 descended through the MVST and c-LVST, respectively. The remaining three were polysynaptic neurons. Histological analysis showed that most of the PC nerve-activated vestibulospinal neurons were located within a specific area in the medial part of the lateral vestibular nucleus and the rostral part of the descending vestibular nucleus. In conclusion, it was suggested that PC nerve-activated vestibulospinal neurons that were located within a focal area of the vestibular nuclei have strong connections with the lower segments of the spinal cord and are related to postural stability that is maintained by the short latency vestibulospinal reflex.     

Otolith-activated vestibulothalamic neurons in cats

The components of the vestibular ascending pathway that transmit otolith information to the thalamus were studied electrophysiologically in anesthetized cats. Thalamic-projecting vestibular neurons (confirmed antidromically) were recorded extracellularly in the various vestibular nuclei. Otolith inputs to these neurons were examined with selective stimulation of the utricular (UT) or the saccular (SAC) nerves. Vestibular nerve branches other than the tested nerve were transected. Of 40 UT-activated vestibulothalamic neurons, 40% (16/40) were activated by UT nerve stimulation with latencies ranging between 0.9-1.4 ms, suggesting they were second-order neurons from the UT nerve. UT-activated vestibulothalamic neurons were recorded in the medial vestibular nucleus (MVN; 24/40), the lateral vestibular nucleus (LVN; 9/40), the descending vestibular nucleus (DVN; 6/40), and the superior vestibular nucleus (SVN; 1/40). Most of the neurons (38/40) were antidromically activated by focal stimulation of the ventral part of the ipsilateral thalamus. Antidromic stimulation of the pontine area revealed that trajectories of the ascending axons (14 of 38 neurons) to the ipsilateral thalamus passed through the pontine reticular formation, ventral to the ascending tract of Deiters (ATD) and the medial longitudinal fasciculus (MLF). Only three SAC-activated vestibulothalamic neurons were encountered in the LVN. All these neurons were second-order neurons from the SAC nerve and were antidromically activated by stimulation of the contralateral thalamus, in marked contrast to the UT-activated vestibulothalamic neurons. Only three UT-activated and two SAC-activated neurons sent descending collaterals to the spinal cord.     

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.     

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.     

Inputs from regularly and irregularly discharging vestibular nerve afferents to secondary neurons in squirrel monkey vestibular nuclei. III. Correlation with vestibulospinal and vestibuloocular output pathways.

1. A previous study measured the relative contributions made by regularly and irregularly discharging afferents to the monosynaptic vestibular nerve (Vi) input of individual secondary neurons located in and around the superior vestibular nucleus of barbiturate-anesthetized squirrel monkeys. Here, the analysis is extended to more caudal regions of the vestibular nuclei, which are a major source of both vestibuloocular and vestibulospinal pathways. As in the previous study, antidromic stimulation techniques are used to classify secondary neurons as oculomotor or spinal projecting. In addition, spinal-projecting neurons are distinguished by their descending pathways, their termination levels in the spinal cord, and their collateral projections to the IIIrd nucleus. 2. Monosynaptic excitatory postsynaptic potentials (EPSPs) were recorded intracellularly from secondary neurons as shocks of increasing strength were applied to Vi. Shocks were normalized in terms of the threshold (T) required to evoke field potentials in the vestibular nuclei. As shown previously, the relative contribution of irregular afferents to the total monosynaptic Vi input of each secondary neuron can be expressed as a %I index, the ratio (x100) of the relative sizes of the EPSPs evoked by shocks of 4 x T and 16 x T. 3. Antidromic stimulation was used to type secondary neurons as 1) medial vestibulospinal tract (MVST) cells projecting to spinal segments C1 or C6; 2) lateral vestibulospinal tract (LVST) cells projecting to C1, C6; or L1; 3) vestibulooculo-collic (VOC) cells projecting both to the IIIrd nucleus and by way of the MVST to C1 or C6; and 4) vestibuloocular (VOR) neurons projecting to the IIIrd nucleus but not to the spinal cord. Most of the neurons were located in the lateral vestibular nucleus (LV), including its dorsal (dLV) and ventral (vLV) divisions, and adjacent parts of the medial (MV) and descending nuclei (DV). Cells receiving quite different proportions of their direct inputs from regular and irregular afferents were intermingled in all regions explored. 4. LVST neurons are restricted to LV and DV and show a somatotopic organization. Those destined for the cervical and thoracic cord come from vLV, from a transition zone between vLV and DV, and to a lesser extent from dLV. Lumbar-projecting neurons are located more dorsally in dLV and more caudally in DV. MVST neurons reside in MV and in the vLV-DV transition zone.(ABSTRACT TRUNCATED AT 400 WORDS)     

Properties of saccular nerve-activated vestibulospinal neurons in cats

Axonal pathways, projection levels, conduction velocities, and locations of the cell bodies of saccular nerve-activated vestibulospinal neurons were studied in decerebrated cats and anesthetized cats, using a collision test of orthodromic and antidromic spikes. The saccular nerve was selectively stimulated by bipolar tungsten electrodes. Three monopolar electrodes were inserted into the left and right lateral vestibulospinal tract (LVST) and medial vestibulospinal tract (MVST) of the C1 segment, to determine the pathway of axons. Three pairs of similar electrodes were positioned bilaterally in the C3–4, T1, and L3 segments to examine projection levels. Another monopolar electrode was placed in the oculomotor nucleus to determine whether saccular nerve-activated vestibulospinal neurons have branches ascending to the oculomotor nucleus. Of 145 vestibular neurons orthodromically activated by stimulation of the saccular nerve, 46 were activated from the C1 segment antidromically. Forty-three were second-order vestibulospinal neurons and 3 were third-order vestibulospinal neurons. Four saccular nerve-activated vestibulospinal neurons were also antidromically activated from the oculomotor nucleus. Sixty-three percent of the saccular nerve-activated vestibulospinal neurons descended through the MVST; one-third of these terminated in the upper cervical segments, one-third reached the lower cervical segments and the remaining one-third reached the upper thoracic segments. Thirty percent of the saccular nerve-activated vestibulospinal neurons descended through the ipsilateral LVST; most of these reached the upper thoracic segments. Seven percent of the saccular nerve-activated vestibulospinal neurons descended through the contralateral vestibulospinal tracts terminating in the upper cervical segments. Most of the saccular nerve-activated vestibulospinal neurons originated in the caudal part of the lateral nucleus and rostral part of the descending nucleus. Received: 8 July 1996 / Accepted: 21 April 1997     

Properties of utricular and saccular nerve-activated vestibulocerebellar neurons in cats

Properties of otolith inputs to vestibulocerebellar neurons were investigated in 14 adult cats. In the vestibular nuclei, we recorded single-unit activities that responded orthodromically after stimulation of the utricular and/or saccular nerves and antidromically after stimulation of the cerebellum (uvula-nodulus and anterior vermis). Descending axonal projections to the spinal cord were also examined by antidromic stimulation of the caudal end of the C1 segment. Forty-seven otolith-activated neurons that projected to the uvula-nodulus were recorded. Thirteen (28%) of the 47 neurons received convergent inputs from the utriculus and sacculus. The remaining 34 (72%) vestibular neurons were non-convergent neurons: 18 (38%) received utricular input alone, and 16 (34%) received saccular input alone. Most (35/47) vestibulocerebellar neurons were located in the descending vestibular nucleus and only one of these projected to the spinal cord. Seven of the 47 vestibulocerebellar neurons were located in the lateral vestibular nucleus and most of these neurons projected to the spinal cord. The remaining neurons were located in group X (two neurons) and the superior vestibular nucleus (three neurons). In a different series of experiments, 37 otolith-activated vestibular neurons were tested to determine whether they projected to the uvula-nodulus and/or the anterior vermis. Nineteen of the 37 neurons projected to the anterior vermis, 13/37 projected to the uvula-nodulus, and 5/37 projected to both. The utricular and/or saccular nerve-activated vestibulocerebellar neurons projected to not only the uvulanodulus, but also to the anterior vermis. In summary, the results of this study showed that vestibular neurons receiving inputs from the utriculus and/or sacculus projected to the cerebellar cortex. This indirect otolith-cerebellar pathway terminated both in the anterior lobe and in the uvula/nodulus.     

Response of pontomedullary reticulospinal neurons to vestibular stimuli in vertical planes. Role in vertical vestibulospinal reflexes of the decerebrate cat.

1. To investigate the neural substrate of vestibulospinal reflexes in decerebrate cats, we studied the responses of pontomedullary reticulospinal neurons to natural stimulation of the labyrinth in vertical planes. Our principal aim was to determine whether reticulospinal neurons that terminate in, or are likely to give off collaterals to, the upper cervical segments had properties similar to those of the vestibulocollic reflex (VCR). 2. Antidromic stimulation was used to determine whether the neurons projected to the neck, lower cervical, thoracic, or lumbar levels. Dynamics of the responses of spontaneously firing neurons were studied with sinusoidal stimuli delivered at 0.05-1 Hz and aligned to the plane of body rotation, that produced maximal modulation of the neuron (response vector orientation). Each neuron was assigned a vestibular input classification of otolith, vertical canal, otolith + canal, or spatial-temporal convergence (STC). 3. We found, in agreement with previous studies, that the largest fraction of pontomedullary reticulospinal neurons projected to the lumbar cord, and that only a small number ended in the neck segments. Neurons projecting to all levels of the spinal cord had similar responses to labyrinth stimulation. 4. Reticulospinal neurons that received only vertical canal inputs were rare (1 of 67 units). Most reticulospinal neurons (48%) received predominant otolith inputs, 18% received otolith + canal input, and only 9% had STC behavior. These data are in sharp contrast to the results of our previous studies of vestibulospinal neurons. A considerable portion of vestibulospinal neurons receives vertical canal input (38%), fewer receive predominantly otolith input (22%), whereas the proportion that have otolith + canal input or STC behavior is similar to our present reticulospinal data. 5. The response vector orientations of our reticulospinal neurons, particularly those with canal inputs (canal, otolith + canal, STC) were predominantly in the roll quadrants. There was no evidence of convergence of inputs from like canals across the midline (e.g., right anterior + left anterior). 6. Two characteristics of the VCR, STC behavior and bilateral input from symmetric vertical canals (in some muscles), cannot be accounted for by the reticulospinal neurons that we studied. Because these characteristics are also not seen in vestibulocollic neurons, they are likely to be the result of the appropriate convergence of vestibular signals in the spinal cord. 7. Pontomedullary reticulospinal neurons seem particularly well suited to play a role in gravity-dependent postural reflexes of neck and limbs.     

Vestibulo-reticular projections in adult lamprey: their role in locomotion

This study describes the anatomical projections from vestibular secondary neurons to reticulospinal neurons in the adult lamprey and the modulation of vestibular inputs during fictive locomotion. Anatomical tracers were applied in the posterior (PRRN) and middle rhombencephalic reticular nuclei as well as to the proximal stumps of cut vestibular nerve branches to identify the neurons projecting to the reticular nuclei that were in close proximity with vestibular primary afferents. Labeled neurons were found in the intermediate (ION) and posterior (PON) octavomotor nuclei, and were more numerous on the side of the injection (around 56-87 and 101-107 for the ION and the PON, respectively). Morphologies varied but cells were mostly round or oval. Axonal projections from the PON formed a dense bundle, whereas those from the ION were less densely packed. Based on their morphology and the distribution of their projections, most vestibulo-reticular neurons were presumed to be vestibulospinal cells. Reticulospinal cells from the PRRN were recorded intracellularly in the in vitro brainstem-spinal cord preparation and large excitatory post-synaptic potentials (EPSPs) were evoked following stimulation of the ipsilateral anterior and the contralateral posterior branches of the vestibular nerves, whereas inhibitory post-synaptic potentials (IPSPs) or smaller EPSPs were elicited by stimulation of the ipsilateral posterior or of the contralateral anterior branches. During fictive locomotion, both the excitatory and the inhibitory responses displayed phasic changes in amplitude such that the amplitude of the EPSPs was minimal when the spinal cord activity switched from the ipsilateral to the contralateral side of the recorded reticulospinal cell. The IPSPs were then of maximal amplitude. We propose that this modulation could serve to reduce the influence of vestibular inputs in response to head movements during locomotion.     

Spatial distribution of the vestibulospinal neurons in the frog vestibular nuclei.

In experiments on the preparation of a frog perfused brain (Rana ridibunda), intracellular potentials were recorded from neurons of the vestibular nuclei following stimulation of the vestibular nerve and the spinal cord. The vestibulospinal neurons were identified on the basis of excitatory postsynaptic potentials evoked by the stimulation of the ipsilateral vestibular nerve and antidromic activation from the stimulation of the cervical and lumbar enlargements of the spinal cord. The cells that could be activated antidromically only by cervical cord stimulation have been designated as C cells, and the cells that could also be activated antidromically as a result of lumbar stimulation have been termed L cells. The average conduction velocity determined for C neurons was 10.67 m/s and for L neurons 15.84 m/s. The ratio of C and L neurons over the vestibular nuclear complex was very similar to each other: 52% C neurons and 48% L neurons. The majority of both types of neurons were localized in the lateral vestibular nucleus (58.6%), to a lesser extent in the descending vestibular nucleus (30.7%) and very little in the medial vestibular nucleus (10.6%). In the lateral vestibular nucleus, C neurons prevailed in the caudal part of the nucleus and L neurons prevailed in the rostral part. By contrast, in the descending and medial vestibular nuclei there was a gradual increase of C and L cells quantitatively from the rostral to the caudal part. Fast and slow cells were detected among the vestibulospinal neurons. The fast neurons of L cells did not prevail greatly over the slow ones, whereas the slow neurons of C cells prevailed comparatively largely over the fast neurons. Thus, it became possible to reconstruct the spatial distribution of the identified vestibulospinal neurons. The results of spatial distribution of C and L vestibulospinal neurons in the frogs failed to conform to definite somatotopy, which is characteristic of mammalian vestibular nuclei.The results of this study have confirmed an earlier assumption that C and L neurons in the frog's vestibular nuclei as a source of vestibulospinal fibers, are scattered separately or more frequently in groups, so that they establish a 'patch-like' somatotopy and do not form a distinctly designed field as in mammals.     

Influence of neck afferents on vestibulospinal neurons

Summary The effects of neck afferent stimulation on vestibulospinal neurons in Deiters' nucleus and in the descending nucleus, and the interaction of cervical and vestibular input, were examined extracellularly in decerebrate, decerebellate cats. Many of the vestibulospinal neurons were identified as having axons in the lateral or medial vestibulospinal tract (LVST or MVST) and as being driven antidromically from C₃ or C₆.Half of the spontaneously active neurons were excited with a latency of 2.5–5.5 ms (early excitation) by stimulation of the contralateral C₂ ganglion. In some neurons early excitation was followed by late excitation (latency > 6 ms), which was in other neurons the only effect seen. Early excitation was due to stimulation of proximal afferents because stimulation of the C₂ dorsal or ventral rami usually produced late excitation only. Early excitation was seen in LVST and MVST neurons terminating between C₃ and C₆ and in those projecting beyond C₆. Neurons with early excitation were scattered throughout Deiters' nucleus and the rostral part of the descending nucleus.In some neurons, mainly in the descending nucleus, the initial effect of contralateral C₂ ganglion stimulation was inhibition. Inhibition could be evoked by stimulation of the ganglion or dorsal rami bilaterally. The axons of all tested inhibited neurons were in the MVST.Thirty-five percent of the population studied received convergence of early excitation and short-latency input from the labyrinth, sometimes from the semicircular canals. There was also convergence between late excitation or inhibition and vestibular input.The influence of neck afferent input on vestibulospinal neurons provides one pathway for this input to the neck and limb segments of the spinal cord. This pathway may be part of the substrate of the tonic neck reflex. In addition, vestibulospinal neurons are one site of interaction between neck and vestibular reflexes.Supported by N.I.H. grant NS 02619     

Input-output relations of lobules I and II of the cerebellar anterior lobe vermis in connexion with neck and vestibulospinal reflexes in the cat

In the anaesthetized cat, lobules I and II of the cerebellar anterior lobe vermis were examined to determine their role in the vestibulospinal and neck-vestibulospinal reflexes with respect to: the somatotopic representation of afferent inputs from labyrinth, neck and tail; and the inhibitory influence on vestibulospinal tract (VST) neurones receiving vestibular and neck afferent inputs. After electrical stimulation of the vestibular nerve and of neck afferents, almost identical responses via mossy fibres were evoked in the lobules, with the prominent response in lobules I and IIa of Larsell. Stimulation of the nerve supplying the dorsal region of the tail induced primarily the mossy fibre response, but also the climbing fibre response, in lobule II. The most responsive areas to tail and neck afferent stimulation did not overlap each other. In the lateral vestibular nucleus, 163 antidromically identified VST neurones were recorded extra- or intracellularly. On the basis of the response pattern to contralateral neck afferent stimulation, they were classified into 3 groups: neurones with excitation (n = 45); neurones with inhibition (n = 71); and neurones with no modulation (n = 47). Stimulation of lobules I-IIa inhibited the activities of 44 VST neurones. Out of them, 41 neurones belonged to the first group. They made up 91% of the group. Twenty-nine of these neurones, i.e. neurones receiving excitatory inputs from the neck and inhibitory inputs from the lobules, received additional excitatory input from the labyrinth. Although lobules I-IIa may be regarded as neck area in the anterior lobe vermis from the viewpoint of sensory input, they did not exert inhibitory influence only exceptionally on vestibulocollic neurones, but predominantly on VST neurones sending their axons to lower thoracic or more caudal segments in the spinal cord. It is suggested from these results that lobules I-IIa have a close relationship with the neck reflex and/or interaction of neck and vestibulospinal reflexes being concerned with the postural adjustment of a rather wide area of the body.     

Convergence of sensory inputs upon projection neurons of somatosensory cortex: Vestibular,neck, head,and forelimb inputs

Summary Cortico-cortical neurons and pyramidal tract (PT) neurons of the cat cerebral cortex were tested for convergent inputs from electrically stimulated vestibular, neck, head and forelimb nerves. Neurons were recorded within forelimb and vestibular projection regions of cortical area 3a. Consideration was given to both suprathreshold and subthreshold inputs. Neither vestibular, neck nor head inputs were detected in the forelimb region of area 3a. In contrast, within the vestibular projection region of area 3a, 43% (6/14) of the cortico-cortical neurons and 63% (24/38) of the PT neurons received excitatory vestibular input. Inputs from the skin of the pinna (greater auricular nerve) were detected only for PT neurons (66%, 25/38). No inputs were detected from afferent nerves supplying the dorsal neck muscles biventer cervicis and complexus. Cortico-cortical and PT neurons receiving vestibular input also received convergent inputs originating from forelimb group I deep and low threshold cutaneous afferent fibers. Further, one half of the PT neurons with vestibular input (12/24) received input from three somatic sources: forelimb group I deep, forelimb low threshold cutaneous and greater auricular (head) nerves. The input connectivities suggest a role for these projection neurons of somatosensory cortex in the coordination of head and forelimb movements. The convergence of vestibular information with somatic input from the forelimb implies that vestibular-influenced neurons of area 3a projecting to the motor cortex or through the pyramidal tract would signal head position or movement with respect to proprioceptive feedback from the limbs.Supported by the Medical Research Council of Canada (MT-7373), the Harry Botterell Foundation for the Neurological Sciences, the Ontario Ministry of Health, the Faculty of Medicine, and the School of Graduate Studies and Research, Queen's University. The visit of P. S. Blum was supported by Queen's QuestRecipient of an Ontario Graduate ScholarshipRecipient of a Medical Research Council of Canada Studentship     

Responses of neurons in the caudal medullary raphe nuclei of the cat to stimulation of the vestibular nerve

Summary In the decerebrate cat, recordings were made from neurons in the caudal medullary raphe nuclei to determine if they responded to electrical stimulation of the vestibular nerve and thus might participate in vestibulo-sympathetic reflexes. Many of these cells projected to the upper thoracic spinal cord. The majority (20/28) of raphespinal neurons with conduction velocities between 1 and 4 m/s received vestibular inputs; 13 of the 20 were inhibited, and 7 were excited. Since many raphespinal neurons with similar slow conduction velocities are involved in the control of sympathetic outflow, as well as in other functions, these cells could potentially relay vestibular signals to sympathetic preganglionic neurons. The onset latency of the vestibular effects was long (median of 15 ms), indicating the inputs were polysynaptic. In addition, 34 of 42 raphespinal neurons with more rapid conduction velocities (6–78 m/s) also received long-latency (median of 10 ms) labyrinthine inputs; 26 were excited and 8 were inhibited. Although little is known about these rapidlyconducting cells, they do not appear to be involved in autonomic control, suggesting that the function of vestibular inputs to raphe neurons is not limited to production of vestibulosympathetic reflexes. One hypothesis is that raphe neurons are also involved in modulating the gain of vestibulocollic and vestibulospinal reflexes; this possibility remains to be tested.     

Electrophysiological analysis of the vestibulospinal reflex pathway of rabbit. I. Classification of tract cells

Summary In anaesthetized rabbits, the medulla was surveyed with recording microelectrodes to identify different types of vestibulospinal tract neurones. Field potentials, unitary extracellular spikes and intracellular potentials were recorded during antidromic stimulation at C₁ and C₆ segments and during orthodromic stimulation through VIIIth nerve. The lateral and medial vestibulospinal tracts (LVST and MVST) were stimulated discriminately with the method developed in Appendix. On the basis of different axonal courses and conduction velocities, three major groups were distinguished for those cells which were activated monosynaptically by the primary vestibular afferents; 1. fast conducting LVST; 2. fast conducting MVST; and 3. slowly conducting MVST. Three other groups were discriminated for those cells which received only a polysynaptic or no action from primary vestibular afferents. These were; 4. fast conducting LVST; 5. slowly conducting LVST and 6. slowly conducting MVST. All of these six types of VST cells were represented within Deiters' nucleus. Only a relatively small number of MVST cells were found in the medial vestibular nucleus.     

Inputs from regularly and irregularly discharging vestibular nerve afferents to secondary neurons in the vestibular nuclei of the squirrel monkey. II. Correlation with output pathways of secondary neurons

1. Intracellular recordings were made from secondary neurons in the vestibular nuclei of barbiturate-anesthetized squirrel monkeys. Monosynaptic excitatory postsynaptic potentials (EPSPs) evoked by stimulation of the ipsilateral vestibular nerve (Vi) were measured. An electrophysiological paradigm, described in the preceding paper (26), was used to determine the proportion of irregularly (I) and regularly (R) discharging Vi afferents making direct connections with individual secondary neurons. The results were expressed as a % I index, an estimate for each neuron of the percentage of the total Vi monosynaptic input that was derived from I afferents. The secondary neurons were also classified as I, R, or M cells, depending on whether they received their direct Vi inputs predominantly from I or R afferents or else from a mixture (M) of both kinds of Vi fibers. The neurons were located in the superior vestibular nucleus (SVN) or in the rostral parts of the medical or lateral (LVN) vestibular nuclei. 2. Antidromic activation or reconstruction of axonal trajectories after intrasomatic injection of horseradish peroxidase (HRP) was used to identify three classes of secondary neurons in terms of their output pathways: 1) cerebellar-projecting (Fl) cells innervating the flocculus (n = 26); 2) rostrally projecting (Oc) cells whose axons ascended toward the oculomotor (IIIrd) nucleus (n = 27); and 3) caudally projecting (Sp) cells with axons descending toward the spinal cord (n = 13). Two additional neurons, out of 21 tested, could be antidromically activated both from the level of the IIIrd nucleus and from the spinal cord. 3. The Vi inputs to the various classes of relay neurons differed. As a class, Oc neurons received the most regular inputs. Sp neurons had more irregular inputs. Fl neurons were heterogeneous with similar numbers of R, M, and I neurons. The mean values (+/- SD) of the % I index for the Oc, Fl, and Sp neurons were 34.7 +/- 24.7, 51.9 +/- 30.4, and 61.8 +/- 18.0%, respectively. Only the Oc neurons had a % I index that was similar to the proportion of I afferents (34%) in the vestibular nerve (cf. Ref. 26). 4. The commissural inputs from the contralateral vestibular nerve (Vc) also differed for the three projection classes. Commissural inhibition was most common in Fl cells: 22/25 (88%) of the neurons had Vc inhibitory postsynaptic potentials (IPSPs) and 1/25 (4%) had a Vc EPSP. In contrast, Vc inputs were only observed in approximately half the Oc and Sp neurons.(ABSTRACT TRUNCATED AT 400 WORDS)