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P K Rose

Publications and source records attributed to P K Rose.

At least 19 recordsLinked to original sources

Distribution of contacts from vestibulospinal axons on the dendrites of splenius motoneurons.

Current descriptions of the organization of synapses on the dendritic trees of spinal motoneurons indicate that the inputs are arranged in several patterns: some are widely distributed; some are distributed to proximal dendrites; others are distributed based on the trajectory of the dendrites. However, the principles governing the organization of synapses on spinal motoneurons remain poorly defined. Our goal was to extend the descriptions of the distribution of synapses, identified by their source, on the dendritic trees of spinal motoneurons. We combined anterograde and intracellular staining techniques in cats to determine the distribution of contacts between excitatory axons from the rostral aspect of the descending vestibular nucleus and the dendrites of motoneurons supplying a dorsal neck muscle, splenius. In five of five motoneurons, the contacts were preferentially distributed on dendrites medial to the soma. This qualitative observation was confirmed by using Monte Carlo methods. The results from this analysis showed that the distribution of contacts can be explained not by the overall distribution of the dendritic membrane area but rather by a systematic innervation of the medial regions of the dendritic trees (P < 0.02). Despite this selectivity, there was no additional bias in the distribution of contacts to proximal vs. distal dendrites. By concentrating excitatory synapses in a restricted region of the dendritic tree, the actions of vestibulospinal connections on neck motoneurons may be increased as a result of a greater probability of activating persistent inward currents on the dendrites.

Animals↗

Effect of nonlinear summation of synaptic currents on the input-output properties of spinal motoneurons.

A single spinal motoneuron receives tens of thousands of synapses. The neurotransmitters released by many of these synapses act on iontotropic receptors and alter the driving potential of neighboring synapses. This interaction introduces an intrinsic nonlinearity in motoneuron input-output properties where the response to two simultaneous inputs is less than the linear sum of the responses to each input alone. Our goal was to determine the impact of this nonlinearity on the current delivered to the soma during activation of predetermined numbers and distributions of excitatory and inhibitory synapses. To accomplish this goal we constructed compartmental models constrained by detailed measurements of the geometry of the dendritic trees of three feline motoneurons. The current "lost" as a result of local changes in driving potential was substantial and resulted in a highly nonlinear relationship between the number of active synapses and the current reaching the soma. Background synaptic activity consisting of a balanced activation of excitatory and inhibitory synapses further decreased the current delivered to the soma, but reduced the nonlinearity with respect to the total number of active excitatory synapses. Unexpectedly, simulations that mimicked experimental measures of nonlinear summation, activation of two sets of excitatory synapses, resulted in nearly linear summation. This result suggests that nonlinear summation can be difficult to detect, despite the substantial "loss" of current arising from nonlinear summation. The magnitude of this "loss" appears to limit motoneuron activity, based solely on activation of iontotropic receptors, to levels that are inadequate to generate functionally meaningful muscle forces.

Animals↗

Relationship between morphoelectrotonic properties of motoneuron dendrites and their trajectory.

The distribution and geometry of the dendritic trees of spinal motoneurons obey several well-established rules. Some of these rules are based on systematic relationships between quantitative geometrical features (e.g., total dendritic length) and the three-dimensional trajectory followed by dendrites from their origin to their termination. Because dendritic geometry partially determines the transmission of current and voltage signals generated by synapses on the dendritic tree, our goal was to compare the efficacy of signal transmission by dendritic trajectories that followed different directions. To achieve this goal, we constructed detailed compartmental models of the dendritic trees of three intracellularly stained biventer cervicis/complexus (BCCM) motoneurons and calculated the electronic properties of 361 dendritic paths. Each trajectory was classified according to its orientation, e.g., rostral, rostral-dorsal-lateral. The attenuation of current and voltage signals en route to the soma was strongly related to trajectory orientation. Trajectories with similar attenuation factors formed functional subunits that were arranged in distinct domains within the ventral horn. Changes in R(m) or R(i) had little effect on which trajectories belonged to each functional subunit. However, differences in the efficacy of signal transmission between subunits increased during high network activity (mimicked by decreases in R(m)). The most efficient subunit delivered two times more current and four times more voltage to the soma than the least efficient subunit. These results indicate that the input-output properties of motoneurons depend on the direction of the path taken by dendrites from their origin at the cell body to their terminals.

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Comparison of the morphological and electrotonic properties of Renshaw cells, Ia inhibitory interneurons, and motoneurons in the cat.

The morphological and electrotonic properties of 4 motoneurons, 8 Ia inhibitory interneurons, and 4 Renshaw cells were compared. The morphological analysis, based on 3-D reconstructions of the cells, revealed that dendrites of motoneurons are longer and more extensively branched. Renshaw cells have dendrites that are shorter and simpler in structure. Dendrites of Ia inhibitory interneurons could be as long as those of motoneurons but the branching structure resembled that of Renshaw cells. Compartmental models were used to determine the electrotonic properties of the paths from each dendritic terminal to the soma. The attenuations of steady-state voltage changes in motoneurons were 3 and 7 times larger than in Ia inhibitory interneurons and Renshaw cells, respectively. The same relative order was observed for current attenuation and electrotonic length. The dendritic input resistances in Renshaw cells were 2 and 4 times larger than in Ia inhibitory interneurons and motoneurons, respectively. The difference in these electrotonic properties increased during higher synaptic activity as modeled by a decrease of Rm. The peak amplitudes of voltage transients at sites of brief, synaptic-like changes in conductance were highly dependent on cell class and were largest in Renshaw cells and smallest in motoneurons. In combination with class-specific differences in the attenuation of transient voltage signals, this led to large differences in the peak amplitudes of somatic voltage transients. Differences in the rise times and half-widths of the voltage transients were observed as well. Thus, based on passive properties, each cell class has a unique set of input/output properties.

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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.

Animals↗

Emergence of axons from distal dendrites of adult mammalian neurons following a permanent axotomy.

The distinctive features of axons and dendrites divide most neurons into two compartments. This polarity is fundamental to the ability of most neurons to integrate synaptic signals and transmit action potentials. It is not known, however, if the polarity of neurons in the adult mammalian nervous system is fixed or plastic. Following axotomy, some distal dendrites of neck motoneurons in the adult cat give rise to unusual processes that, at a light microscopic level, resemble axons (Rose, P.K. & Odlozinski, M., J. Comp. Neurol., 1998, 390, 392). The goal of the present experiments was to characterize these unusual processes using well-established ultrastructural and molecular criteria that differentiate dendrites and axons. These processes were immunoreactive for growth-associated protein-43 (GAP-43), a protein that is normally confined to axons. In contrast, immunoreactivity for a protein that is widely used as a marker for dendrites, microtubule-associated protein (MAP)-2a/b, could not be detected in the unusual distal arborizations. At the electron microscopic level, unusual distal processes contained dense collections of neurofilaments and were frequently myelinated. These molecular and structural characteristics are typical of axons and suggest that the polarity of adult neurons in the mammalian nervous system can be disrupted by axotomy. If this transformation in neuronal polarity is common to other types of neurons, axon-like processes emerging from distal dendrites may represent a mechanism for replacing connections lost due to injury. Alternatively, the connections formed by these axons may be aberrant and therefore maladaptive.

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Projections from the lateral vestibular nucleus to the upper cervical spinal cord of the cat: A correlative light and electron microscopic study of axon terminals stained with PHA-L.

Vestibulospinal axon collaterals in C1 and C2 were stained following injections of Phaseolus vulgaris leucoagglutinin (PHA-L) into the lateral vestibular nucleus (LVN). The distribution and geometry of collaterals within three regions of the ventral horn were determined at the light microscopic level. These processes were subsequently examined at the electron microscopic level to define the relationship between their ultrastructural characteristics and their geometry and location. All round or elliptical varicosities, whose diameters exceeded the diameter of the adjacent axon shaft by a factor of two, as measured at the light microscopic level, contained synaptic vesicles and contacted dendrites or somata. These varicosities accounted for 82% of labelled axon terminals found at the electron microscopic level. Thus, axon terminals stained with PHA-L can be identified reliably at the light microscopic level, but synaptic density will be slightly underestimated. One-hundred and thirty-eight axon terminals were classified as excitatory or inhibitory on the basis of well-established morphological criteria (e.g., vesicle shape). Placed in the context of previous physiological observations describing the excitatory or inhibitory actions of medial and lateral vestibulospinal tract (MVST and LVST) neurons, our results suggest that projections from the LVN to the ipsilateral ventral horn originate primarily from the LVST. These connections are excitatory. Ipsilateral connections via the MVST are inhibitory and are largely confined to a region near the border of laminae VII and VIII. Most axon terminals in the contralateral ventral horn were inhibitory. This result indicates that the LVN is the source of a specific subset of crossed MVST axons with inputs from the posterior semicircular canal.

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Cortical influences on the vestibular nuclei of the cat.

Our goal was to study potential substrates for cortical modulation of vestibular reflexes in the cat. In initial experiments, injections of wheat-germ-agglutinate-horseradish-peroxidase into Deiters' nucleus and the rostral descending nucleus revealed bilateral colonies of retrogradely filled neurons in cortical areas 6, 2, and 3a (about 60 cells per colony). In cats anesthetized with chloralose-urethane, we stimulated areas 2 and 3a with trains of pulses while recording from ipsilateral vestibular-nucleus neurons, which were characterized by their responses to sinusoidal tilts and tested for the presence of antidromic responses to stimulation of the upper cervical cord. A majority of the neurons was affected by cortical stimulation, showing either facilitation, inhibition, or a mixture of the two. Stimulation in area 2 was more effective than stimulation in area 3a. Despite the anatomic presence of direct cortico-vestibular projections, properties of facilitation and inhibition suggest that both were evoked by polysynaptic pathways. Cortical effects were broadly distributed to vestibular neurons without regard to responses of these neurons to sinusoidal tilts. There was no significant difference between effects on lateral and medial vestibulospinal tract neurons, but, as a group, vestibulospinal neurons were much more likely to be affected by cortical stimulation than neurons not antidromically activated from the C2 segment. We conclude that, by their influence on vestibulospinal neurons, neurons in cortical areas 2 and 3a should be able to modulate, in behaving animals, vestibular reflexes acting on the neck and limbs.

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Expansion of the dendritic tree of motoneurons innervating neck muscles of the adult cat after permanent axotomy.

The morphologic characteristics of neck motoneurons with intact axons were compared with those of neck motoneurons that had been permanently axotomized for 11 to 17 weeks. Motoneurons were identified antidromically, intracellularly stained with horseradish peroxidase (HRP) and examined after reconstructions of their entire dendritic tree. Axotomized motoneurons differed qualitatively and quantitatively from motoneurons with intact axons. The distal branches of axotomized motoneurons exhibited two novel features: some gave rise to tangled appendages that exhibited growth cone-like specializations resembling lamellipodia and filopodia; others followed a meandering path and had unusually large diameters. These branches showed a discontinuous pattern of staining that was similar to the appearance of myelinated axons stained intra-axonally with HRP. A quantitative analysis of the dendritic trees of 13 completely reconstructed dendritic trees (five axotomized motoneurons and eight motoneurons with intact axons) showed that total dendritic surface area, total dendritic length, and total number of branches increased 38, 34, and 215%, respectively, after axotomy. These measurements were confirmed by comparing the sizes of a larger number of motoneurons (16 axotomized and 21 intact), calculated on the basis of correlations between dendritic tree size and proximal dendritic diameter. We conclude, therefore, that neck motoneurons, in contrast to other types of motoneurons, expand their dendritic trees after axotomy. It is suggested that this expansion is a consequence of two mechanisms: one involves dendritic growth, possibly leading to new synaptic connections; the other causes a conversion of some dendrites into axons.

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Contribution of voltage-dependent potassium channels to the somatic shunt in neck motoneurons of the cat.

The specific membrane resistivity of motoneurons at or near the soma (Rms) is much lower than the specific membrane resistivity of the dendritic tree (Rmd). The goal of the present experiments was to investigate the contribution of tonically active voltage-dependent potassium channels at or near the soma to the low Rms. These channels were blocked with the use of intracellular injections of cesium. Input resistance (RN), Rms/Rmd, a conductance representing voltage-dependent potassium channels on the soma, GK, and a conductance attributed to damage caused by electrode impalement, GDa, were estimated from voltage responses to a step of current. The effect of intracellular injections of cesium on electrotonic structure was determined with the use of two strategies: 1) a population analysis that compared data from two groups of motoneurons, those recorded with potassium acetate electrodes (control group) and those recorded with cesium acetate electrodes after the motoneurons were loaded with cesium; and 2) an analysis of changes in electrotonic structure that occurred over the course of multiple injections of cesium or during similar periods of time in control cells. RN of control and cesium-loaded motoneurons was similar. Over the course of the recordings, RN of control cells usually increased and this increase was associated with a hyperpolarization. In contrast, increases in RN after successive injections of cesium were closely linked to a depolarization. At corresponding membrane potentials, Rms/Rmd of cesium-loaded motoneurons was greater than Rms/Rmd of control motoneurons. Over the course of cesium injections, Rms/Rmd increased and the membrane potential depolarized. In contrast, increases in Rms/Rmd observed during the course of recordings from control cells were associated with a hyperpolarization. Compared with control cells at corresponding membrane potentials, GK was less in cesium-loaded cells. Increases in GK that occurred over the course of recordings in control cells were closely coupled to a depolarization. In cesium-loaded cells, GK usually decreased as the cell depolarized during the injections. In control cells, increases in GDa that occurred during the recording period were closely coupled to a depolarization. In contrast, changes in GDa that occurred during cesium injections were not related to the change in membrane potential and did not explain the corresponding changes in Rms/Rmd and membrane potential. The results of this study indicate that voltage-dependent potassium channels contribute to the somatic shunt (low Rms) in neck motoneurons and provide a voltage-dependent mechanism for the dynamic regulation of motoneuron electrotonic properties.

Action Potentials↗

Morphology of single vestibulospinal collaterals in the upper cervical spinal cord of the cat: III collaterals originating from axons in the ventral funiculus ipsilateral to their cells of origin.

Some vestibulospinal pathways are composed of a homogeneous collection of axons with similar intraspinal collaterals. Other pathways contain axons whose collaterals vary in terms of shape, distribution, and complexity. The purpose of the present study was to extend the study of homogeneity versus heterogeneity of vestibulospinal axons to vestibulospinal axons that travel in the ventral funiculus ipsilateral to their cells of origin. Collaterals of these axons were stained following extracellular injections of Phaseolus vulgaris-leucoagglutinin in rostral parts of the medial and descending vestibular nuclei. All collaterals found in C2 and C3 were reconstructed. Collaterals arising from small diameter (0.5 to 2.9 microns) axons usually consisted of a single main branch with short side branches. The termination zones of most of these collaterals formed a narrow path in lamina VIII, but the location of this pathway was highly variable. Collaterals arising from large-diameter (3.0-6.1 microns) axons were usually more complex and consisted of many branches with en passant and terminal boutons that were located in motoneuron nuclei as well as laminae VIII and VII. Despite a relationship between termination zone and the position of the parent axon in the ventral funiculus, the variability in collaterals from large-diameter axons precluded a simple classification scheme. These results demonstrate that diversity, instead of homogeneity, is a characteristic feature of vestibulospinal axons that originate from the medial and descending vestibular nuclei and travel in the ipsilateral ventral funiculus. This pathway is therefore composed of multiple anatomical subunits that, as individuals, may selectively coordinate the activity of specific combinations of interneurons and motoneurons.

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Innervation of motoneurons based on dendritic orientation.

1. The distribution of terminals from vestibulospinal (VS) axons on the dendritic trees of neck motoneurons was determined by combining the anterograde transport of Phaseolus Vulgaris Leucoagglutinin (PHA-L) with intracellular staining techniques and three-dimensional reconstruction methods. 2. Contacts between VS axon terminals and dendrites were arranged in a nonuniform pattern that depended on the orientation (i.e., rostro-caudal vs. dorsolateral) of the dendrites. 3. This innervation pattern satisfies a critical structural condition necessary for selective nonlinear interactions between pairs of simultaneously active inputs to motoneurons.

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Organization of single motor units in feline sartorius.

1. We depleted single motor units in feline sartorius muscles of glycogen by stimulating their motoneurons intracellularly. We mapped the intramuscular distribution of depleted fibers by inspecting histological cross-sections throughout the length of sartorius. 2. We selected ten depleted motor units for detailed study and quantitative analysis. Nine motor units were located in the anterior head of sartorius. One was located in a muscle whose distal half appeared to have been damaged some time before the acute experiment. A single motor unit was located in the medial head of sartorius. 3. Five motor units were composed of fast-twitch glycolytic (FG) muscle fibers, two of fast-twitch oxidative glycolytic (FOG) muscle fibers, and three of slow-twitch oxidative (SO) muscle fibers. Estimates of the numbers of depleted fibers in motor units of anterior sartorius indicated that FG motor units were larger (mean 566 fibers) than FOG and SO motor units (SO mean 190, FOG mean 156 fibers). The SO motor unit in the damaged muscle had 550 fibers. One motor unit depleted in the medial head of sartorius had 270 fibers with FG profiles. 4. Muscle fibers belonging to each anterior motor unit were never distributed throughout the whole cross-section of anterior sartorius at any proximodistal level. Furthermore, fibers were distributed nonuniformly along the proximodistal axis of the muscle. In most muscles at least a few depleted fibers were found at all proximodistal levels. However, in one normal muscle and the damaged muscle, depleted fibers were confined to the proximal end. 5. The fibers in the medial motor unit were confined to a strip that did not extend across the whole cross-section of the muscle head. Fibers within this strip were scattered quite evenly from origin to insertion. This medial FG motor unit occupied a smaller territory and contained fewer fibers than anterior motor units of the same histochemical type. 6. These results show that sartorius motor units are not distributed uniformly in the mediolateral plane; those in anterior sartorius were distributed asymmetrically in the proximodistal axis as well. This finding has important functional implications for the way in which we model force development and transmission in sartorius and other long muscles.

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Facial input to neck motoneurons: trigemino-cervical reflexes in the conscious and anaesthetised cat.

Cutaneous facial inputs influencing head movement were examined in the conscious and anaesthetised cat. EMG recordings were made in neck muscles of conscious, unrestrained cats in which an unexpected light cutaneous stimulus was applied to the glabrous skin of the planum nasale (PN). These observations established that head aversion movements were associated with synchronised activation of both deep and superficial dorsal neck muscles. In anaesthetised cats in which activity in the motoneurons of the large dorsal neck muscles was examined, mechanical stimulation of the PN or electrical stimulation of the infraorbital nerve (ION) produced a short latency, reflex activation. The reflex could be elicited by excitation of low threshold, rapidly conducting fibres in the ION. Intracellular recording from neck motoneurons showed that there is a short latency, probably disynaptic, excitatory pathway from low threshold nerves in the ION to neck motoneurons, but discharge of neck motoneurons occurred several milliseconds later, presumably as a result of activity in a longer multisynaptic pathway.

Anesthesia↗

Morphology of single vestibulospinal collaterals in the upper cervical spinal cord of the cat: I. Collaterals originating from axons in the ventromedial funiculus contralateral to their cells of origin.

Vestibulospinal neurons in the medial and descending vestibular nuclei have widespread bilateral terminations in the upper cervical spinal cord. These terminations arise from axons travelling in several funiculi, including the ventromedial, ventrolateral, lateral, and dorsolateral funiculi in addition to the dorsal columns. The purpose of the present study was to examine the morphology of single vestibulospinal collaterals which terminate in the upper cervical spinal cord and which originate from axons located in one of these funicular pathways, the ventromedial funiculus, contralateral (cVMF) to their cells of origin in the vestibular nuclei. The 32 collaterals described were selected from two separate sets of experiments which took advantage of different techniques. Nineteen of the collaterals were labelled following Phaseolus vulgaris leucoagglutinin (PHA-L) injections into the medial vestibular nucleus and medial regions of the descending vestibular nucleus. The remaining 13 collaterals originated from physiologically identified vestibulospinal axons that were stained after intra-axonal injections of horseradish peroxidase (HRP). The combined projection of all cVMF axon collaterals spread from laminae V to IX, and included the central cervical nucleus. There was a high degree of variability in the pattern of terminations of individual collaterals. This variability was more pronounced among PHA-L-labelled collaterals than HRP-labelled collaterals whose terminations were restricted to laminae VIII and IX. Some PHA-L-labelled collaterals had terminations which were focused within a single lamina, whereas others had termination zones spanning as many as four laminae. The differences between collaterals were compounded when the characteristics of branching patterns were considered. Some collaterals which occupied similar termination zones had different branching structures.(ABSTRACT TRUNCATED AT 250 WORDS)

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Morphology of single vestibulospinal collaterals in the upper cervical spinal cord of the cat. II. Collaterals originating from axons outside the ventral funiculi.

Recent studies have shown that vestibulospinal axons reach the upper cervical spinal cord of the cat via several different funicular routes. The purpose of this study was to describe the projections of those axons travelling outside the well-recognized pathways in the ventral funiculi. These axons are located in the dorsal columns, dorsolateral funiculi, and lateral funiculi. Collaterals of these axons were stained following extracellular injections of Phaseolus vulgaris leucoagglutinin in the medial and descending vestibular nuclei. The trajectories of individual collaterals were reconstructed from serial histological sections. Collaterals arising from axons in the same funiculus usually had the same characteristic appearance. Axons in the lateral funiculi, ipsilateral or contralateral to their cells of origin, gave rise to collaterals that had a simple structure and usually followed a horizontal trajectory across laminae VII and VIII. The boutons of these collaterals were distributed throughout the mediolateral extent of laminae VI and VII and the dorsal half of lamina VIII. In contrast, axons in the dorsolateral funiculi, ipsilateral or contralateral to their cells of origin, terminated primarily in laminae IV and V. Many collaterals of these axons projected either rostrally or caudally and had a narrow mediolateral distribution. The combined distribution of boutons from collaterals originating from axons in the dorsal columns included the dorsal horn and intermediate zone. Although these collaterals were less common and formed a heterogeneous group, they were easily distinguished from collaterals originating from axons travelling in other funiculi. These results indicate that vestibulospinal axons travelling outside the ventral funiculi comprise several distinct systems. Each system travels by a different funicular route and is distinguished by differences in collateral morphology and termination zones.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Connections from the lateral vestibular nucleus to the upper cervical spinal cord of the cat: a study with the anterograde tracer PHA-L.

The projections of neurons in the lateral vestibular nucleus (LVN) to the upper cervical spinal cord of the cat were investigated by means of the anterograde tracer Phaseolus vulgaris leucoagglutinin (PHA-L). At the junction of C1 and C2, axons were distributed bilaterally in the ventromedial funiculi, and ipsilaterally in the ventrolateral and lateral funiculi. The majority of boutons were found ipsilateral to the injection sites and most of these boutons were found at the base of the ventral horn and throughout the medial two-thirds of lamina VIII. A more modest termination zone was found along the ventral border of lamina VII and a small number of boutons were scattered in the dorsal horn. Contralateral termination zones were similar to the ipsilateral projections. There were significant changes in the distribution of vestibulospinal axons and density of boutons at the junction of C3 and C4. At this level, most vestibulospinal axons travelled ipsilaterally and were found along the medial border of the ventromedial funiculus and the ventral margin of the ventrolateral funiculus. The overall distribution of boutons near the border of C3 and C4 was similar to the pattern seen at the junction of C1 and C2. However, bouton density fell by a factor of three. Large zones of the grey matter were devoid of boutons in individual experiments. These results demonstrate that the projections of neurons in the LVN to the upper cervical spinal cord are densest in the regions containing motoneurons supplying suboccipital muscles. This result suggests that monosynaptic connections to those motoneurons may be an important part of the neural circuitry responsible for vestibulocollic reflexes. However, the large number of boutons found in regions dorsal to motoneuron nuclei in all upper cervical segments indicates that the primary path from vestibulospinal axons to neck motoneurons may be indirect and involve relays via spinal interneurons.

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