Influence of surgical plugging on horizontal semicircular canal mechanics and afferent response dynamics.

Mechanical occlusion of one or more of the semicircular canals is a surgical procedure performed clinically to treat certain vestibular disorders and used experimentally to assess individual contributions of separate canals and/or otoliths to vestibular neural pathways. The present experiments were designed to determine if semicircular canal afferent nerve modulation to angular head acceleration is blocked by occlusion of the endolymphatic duct, and if not, what mechanism(s) might account for a persistent afferent response. The perilymphatic space was opened to gain acute access to the horizontal canal (HC) in the oyster toadfish, Opsanus tau. Firing rate responses of HC afferents to sinusoidal whole-body rotation were recorded in the unoccluded control condition, during the process of duct occlusion, and in the plugged condition. The results show that complete occlusion of the duct did not block horizontal canal sensitivity; individual afferents often exhibited a robust firing rate modulation in response to whole-body rotation in the plugged condition. At high stimulus frequencies (about >8 Hz) the average sensitivity (afferent gain; spikes/s per degrees /s of head velocity) in the plugged condition was nearly equal to that observed for unoccluded controls in the same animals. At low stimulus frequencies (about <0.1 Hz), the average sensitivity in the plugged condition was attenuated by more than two orders of magnitude relative to unoccluded controls. The peak afferent firing rate for sinusoidal stimuli was phase advanced approximately 90 degrees in plugged canals relative to their control counterparts for stimulus frequencies approximately 0.1-2 Hz. Data indicate that afferents normally sensitive to angular velocity in the control condition became sensitive to angular acceleration in the plugged condition, whereas afferents sensitive to angular acceleration in the control condition became sensitive to the derivative of acceleration or angular jerk in the plugged condition. At higher frequencies (>8 Hz), the phase of afferents in the plugged condition became nearly equal, on average, to that observed in controls. A three-dimensional biomechanical model of the HC was developed to interpret the residual response in the plugged condition. Labyrinthine fluids were modeled as incompressible and Newtonian; the membranous duct, osseous canal and temporal bone were modeled as visco-elastic materials. The predicted attenuation and phase shift in cupular responses were in close agreement with the observed changes in afferent response dynamics after canal plugging. The model attributes the response of plugged canals to labyrinthine fluid pressure gradients that lead to membranous duct deformation, a spatial redistribution of labyrinthine fluids and cupular displacement. Validity of the model was established through its ability to predict: the relationship between plugged canal responses and unoccluded controls (present study), the relationship between afferent responses recorded during mechanical indentation of the membranous duct and physiological head rotation, the magnitude and phase of endolymphatic pressure generated during HC duct indentation, and previous model results for cupular gain and phase in the rigid-duct case. The same model was adjusted to conform to the morphology of the squirrel monkey and of the human to investigate the possible influence of canal plugging in primates. Membranous duct stiffness and perilymphatic cavity stiffness were identified as the most salient model parameters. Simulations indicate that canal plugging may be the most effective in relatively small species having small labyrinths, stiff round windows, and stiff bony perilymphatic enclosures.     

Afferent responses during experimentally induced semicircular canalithiasis

Benign paroxysmal positional vertigo (BPPV) is a common vestibular disorder that results in brief periods of vertigo and nystagmus, when the head is tipped relative to gravity. Symptoms are commonly attributed to the pathological presence of heavy calcium carbonate particles within the lumen of the semicircular canal(s)-a condition termed canalithiasis. In the present work, we induced canalithiasis in an animal model (oyster toadfish, Opsanus tau) by introducing heavy glass microbeads into the lumen of the lateral semicircular canal. Bead movement under the action of gravity and canal afferent nerve discharge were recorded in vivo. When the head was oriented nose-down, beads moved toward the nose and the lateral canal afferent discharge rate increased. Afferents that normally encoded angular velocity during oscillatory head rotations responded with tonic increases in the discharge rate during gravity-dependent bead movement. Other afferents, such as the units that rapidly adapt to a step increase in angular head velocity, responded with an initial increase in discharge rate followed by a period of adaptation. Afferent responses occurred in the complete absence of head movement and quantify the pathological inputs to the brain that arise from canalithiasis. The magnitude and time course of the responses reported here are sufficient to explain the symptoms of BPPV.     

Dynamics of vestibular neurons during rotational motion in alert rhesus monkeys

The temporal processing in the encoding of head rotation was investigated by comparing the dynamics of vestibular nuclei neurons with those of the regularly and irregularly firing semicircular canal afferents in alert rhesus monkeys. During earth-vertical axis rotations, neurons without eye movement sensitivity differed in their response dynamics from both regularly and irregularly firing semicircular canal afferents. At high frequencies, central responses increased in sensitivity and maintained phase leads of nearly 30° relative to head velocity. These persistent high-frequency phase leads resembled those of irregularly firing (but not regularly firing) semicircular canal afferents. However, at low frequencies, central responses exhibited significantly smaller phase leads than those of irregularly firing semicircular canal afferents, and dynamics resembled more those of the regularly firing afferents. The response dynamics of central non-eye movement cells were significantly different from those of position-vestibular-pause and eye-head neurons (collectively referred to as eye movement cells). In contrast to the persistent phase leads of non-eye movement neurons, all eye movement cells modulated closely in phase with head velocity at all frequencies down to 0.05 Hz during visual suppression tasks. Vertical canal non-eye movement neurons that were insensitive to both translations and static head tilts led head velocity by approximately 5–30° during high-frequency earth-horizontal axis rotations. Unlike the earth-vertical axis responses that led head velocity at low frequencies by as much as 20–40°, vertical canal neurons only slightly led or even lagged behind head velocity during low-frequency earth-horizontal axis rotations. Posterior canal central non-eye movement cells lagged behind head velocity significantly more than anterior canal neurons. These frequency dependencies of central vestibular neurons in comparison with those of the afferents suggest that both low- and high-pass filtering might be necessary to convert primary semicircular canal afferent response dynamics to central neuron ones.     

Eye movement responses to combined linear and angular head movement

Summary Lateral eye movements evoked by linear head motion were evaluated in human subjects by subtracting the eye movement responses to headcentred angular oscillation in the dark, about a vertical axis, from the responses evoked by similar oscillation with the head displaced 30 cm eccentrically from the axis. The centred oscillation gave a purely angular stimulus whereas the eccentric oscillation gave an additional tangential linear acceleration acting laterally to the head. The stimuli used were relatively unpredictable, enveloped sinewaves at 0.02 to 1.2 Hz, 60°/s peak angular velocity, 0.004 to 0.24 g peak tangential acceleration, and subjects were either given no instructions or were told to imagine fixating on targets at 60 cm or 5 m distance. Eye movements of significantly higher velocity were evoked in the eccentric position, particularly at the higher frequencies and when subjects imagined near targets. The increase in velocity of eye movement was attributed to the linear stimulus and probably derives from stimulation of the otolith organs. The frequency response of the gain (°/s/g) of these movements gave an approximate slope of –1, indicating that the eye velocity bears a constant proportionality to linear head velocity. The findings are in accord with the theoretical prediction that eye movements compensating for linear head motion should only be required for viewing near targets. These otolithic influences on eye movements could either the mediated by a direct otolith-ocular reflex which is subservient to viewing conditions or, alternatively, the otolith signals may modify the activity of other oculomotor mechanisms.A. M. Bronstein was supported by The Brain Research Trust     

The response of primary horizontal semicircular canal neurons in the rat and guinea pig to angular acceleration

Summary In rats and guinea pigs, primary afferent neurons from the horizontal semicircular canal were divided into two categories, regular and irregular, on the basis of the regularity of their resting activity. Regular neurons tend to have higher average resting rates than irregular neurons and in response to a constant angular acceleration stimulus of 16.7 deg/s² regular neurons tended to have lower sensitivity and longer time constants than irregular cells. Some irregular neurons are more sensitive to incremental accelerations than to decremental accelerations of the same magnitude, whereas regular neurons tend to show symmetrical sensitivity.In response to sinusoidal angular acceleration stimuli (fixed frequencies) in the range 0.01–1.5 Hz, cells which fired regularly at rest tended to have smaller gain and longer phase lag re acceleration at most frequencies than irregular cells. Transfer functions were obtained for averaged data for regular and irregular neurons separately in both species.In both species there is evidence of systematic variation between neurons within each category, and this systematic variation is obscured by averaging across neurons.     

Velocity- and acceleration-sensitive units in the trunk lateral line of the trout.

1. The two main types of lateral line organs of lower vertebrates are the superficial neuromasts (SN), with a cupula that protrudes in the surrounding water, and the canal neuromasts (CN), located in the lateral line canal. The scales of the trunk lateral line canal of fish contain SNs as well as CNs. In this study, we examine whether there exist two functional classes of afferent fibers in the trunk lateral line nerve of the rainbow trout that can be attributed to the SNs and CNs. 2. The response properties of the afferent fibers in the trunk lateral line nerve have been determined during stimulation with sinusoidally varying water motion generated by a small vibrating sphere. Linear frequency response analysis revealed the presence of two distinct populations of afferent fibers in the lateral line nerve. The fibers belonging to the two populations showed significant differences in the frequency at which the sensitivity was maximal, the low-frequency response slope and the low-frequency asymptotic phase angle. 3. One population of fibers has a maximum sensitivity at 36 +/- 13 (SD) Hz (n = 22) and responds up to this frequency to water velocity. The low-frequency slope of the frequency response of these fibers was 20 +/- 3 (SD) dB/decade and the low-frequency phase lead was 121 +/- 11 degrees (mean +/- SD), both with respect to sphere displacement. The fibers of the other population have a maximum sensitivity at 93 +/- 14 (SD) Hz (n = 12) and respond up to this frequency to water acceleration. The low-frequency slope of these fibers was 35 +/- 5 (SD) dB/decade, and the low-frequency phase lead was 188 +/- 13 degrees (mean +/- SD). 4. Analysis of the stochastic properties of the spontaneous activity of both types of fibers revealed that the mean firing rate of the fibers responding to water velocity (26 +/- 12 spikes/s, mean +/- SD; n = 22) was significantly higher than that of the fibers responding to acceleration (36 +/- 11 spikes/s, mean +/- SD; n = 12). The other statistical properties of the spontaneous activity were found to be indistinguishable. 5. From comparison of the results with the available quantitative data on frequency responses of lateral line organs in other species, it has been concluded that the fibers responding (< or = 40 Hz) to water velocity innervate SNs and that the fibers responding (< or = 90 Hz) to water acceleration innervate CNs.(ABSTRACT TRUNCATED AT 400 WORDS)     

Neural transduction in Xenopus laevis lateral line system

1. The process of neural excitation in hair cell systems was studied in an in vitro preparation of the Xenopus laevis (African clawed toad) lateral line organ. A specially designed stimulus chamber was used to apply accurately controlled pressure, water movement, or electrical stimuli, and to record the neural responses of the two afferent fibers innervating each organ or stitch. The objective of the study was to determine the characteristics of the neural responses to these stimuli, and thus gain insight into the transduction process. 2. A sustained deflection of the hair cell cilia due to a constant flow of water past the capula resulted in a maintained change in the mean firing rate (MFR) of the afferent fibers. The data also demonstrated that the neural response was proportional to the velocity of the water flow and indicated that both deflection and movement of the cilia were the effective physiological stimuli for this hair cell system. 3. The preparations responded to sinusoidal water movements (past the capula) over the entire frequency range of the stimulus chamber, 0.1-130 Hz, and were most sensitive between 10 and 40 Hz. The variation of the MFR and the percent modulation indicated that the average dynamic range of each organ was 23.5 dB. 4. The thresholds, if any, for sustained pressure changes and for sinusoidal pressure variations in the absence of water movements were very high. Due to the limitations of the stimulus chamber it was not possible to generate pressure stimuli of sufficient magnitude to elicit a neural response without also generating suprathreshold water-movement stimuli. Sustained pressures had no detectable effect on the neural response to water-movement stimuli. 5. The preparations were very sensitive to electrical potentials applied across the toad skin on which the hair cells were located. Potentials which made the ciliated surfaces of the hair cells positive with respect to their bases increased the MFR of the fibers, whereas negative potentials decreased it. The responses to sinusoidal electrical stimuli were similar to responses to water-movement stimuli with respect to frequency and dynamic ranges. Thresholds as low as 100 muV peak to peak (p-p) for 16-Hz stimuli were found. 6. The characteristics of the neural responses to electrical stimulation as well as supporting data obtained from the studies of the effects of anoxia on the evoked responses indicate that the electrical stimulus acts on the hair cells or on the synapses, rather than directly on the nerve fibers. This finding suggests that receptor potentials or their associated currents play an important role in the process of neural excitation in hair cell systems.     

Spatial orientation of semicircular canals and afferent sensitivity vectors in pigeons

Rotational head motion in vertebrates is detected by the semicircular canal system, whose innervating primary afferent fibers carry information about movement in specific head planes. The semicircular canals have been qualitatively examined over a number of years, and the canal planes have been quantitatively characterized in several animal species. The present study first determined the geometric relationship between individual semicircular canals and between the canals and the stereotactic head planes in pigeons. Stereotactic measurements of multiple points along the circumference of the bony canals were taken, and the measured points fitted with a three-dimensional planar surface. Direction normals to the plane's surface were calculated and used to define angles between semicircular canal pairs. Because of the unusual shape of the anterior semicircular canals in pigeons, two planes, a major and a minor, were fitted to the canal's course. Calculated angle values for all canals indicated that the horizontal and posterior semicircular canals are nearly orthogonal, but the anterior canals have substantial deviations from orthogonality with other canal planes. Next, the responses of the afferent fibers that innervate each of the semicircular canals to 0.5 Hz sinusoidal rotation about an earth-vertical axis were obtained. The head orientation relative to the rotation axis was systematically varied so that directions of maximum sensitivity for each canal afferent could be determined. These sensitivity vectors were then compared with the canal plane direction normals. The afferents that innervated specific semicircular canals formed homogeneous clusters of sensitivity vectors in different head planes. The horizontal and posterior afferents had average sensitivity vectors that were largely coincident with the innervated canal plane direction normals. Anterior canal afferents, however, appeared to synthesize contributions from the major and minor plane components of the bony canal structure to produce a resultant sensitivity vector that was positioned between the canal planes. Calculated angles between the average canal afferent sensitivity vectors revealed that direction orthogonality is preserved at the afferent signal level, even though deviations from canal plane orthogonality exist.     

Cross-axis adaptation improves 3D vestibulo-ocular reflex alignment during chronic stimulation via a head-mounted multichannel vestibular prosthesis

By sensing three-dimensional (3D) head rotation and electrically stimulating the three ampullary branches of a vestibular nerve to encode head angular velocity, a multichannel vestibular prosthesis (MVP) can restore vestibular sensation to individuals disabled by loss of vestibular hair cell function. However, current spread to afferent fibers innervating non-targeted canals and otolith end organs can distort the vestibular nerve activation pattern, causing misalignment between the perceived and actual axis of head rotation. We hypothesized that over time, central neural mechanisms can adapt to correct this misalignment. To test this, we rendered five chinchillas vestibular deficient via bilateral gentamicin treatment and unilaterally implanted them with a head-mounted MVP. Comparison of 3D angular vestibulo-ocular reflex (aVOR) responses during 2 Hz, 50°/s peak horizontal sinusoidal head rotations in darkness on the first, third, and seventh days of continual MVP use revealed that eye responses about the intended axis remained stable (at about 70% of the normal gain) while misalignment improved significantly by the end of 1 week of prosthetic stimulation. A comparable time course of improvement was also observed for head rotations about the other two semicircular canal axes and at every stimulus frequency examined (0.2-5 Hz). In addition, the extent of disconjugacy between the two eyes progressively improved during the same time window. These results indicate that the central nervous system rapidly adapts to multichannel prosthetic vestibular stimulation to markedly improve 3D aVOR alignment within the first week after activation. Similar adaptive improvements are likely to occur in other species, including humans.     

Quantitative aspects of responses in trigeminal relay neurones and interneurones following mechanical stimulation of sinus hairs and skin in the cat

1. Stimulus-response relationships in discharges of trigeminal relay- and interneurones were investigated in the barbiturate anaesthetized cat using controlled sinus hair or skin displacements.2. In comparison with discharges in slowly adapting primary afferent fibres the responses in all higher order neurones were considerably reduced in firing rate and often revealed modifications suggesting the interaction of mechanisms actively modulating the afferent input.3. In relay neurones with or without a tonic discharge component the ;dynamic on' response during a trapezoidal displacement of sinus hairs was found to be determined entirely or predominantly by the movement velocity and to be independent of the deflexion angle of a stimulus. In contrast, the static response in tonic relay neurones was determined by both the movement velocity and the displacement amplitude.4. Spatial summation of afferent input caused either only quantitative changes in the responses of relay neurones leaving the general discharge properties unaltered or caused both qualitative and quantitative changes in the responses.5. Interneurones consisted of two functional groups. In about 25% of them the responses were not or only slightly dependent on the intensity of the applied stimulus, often burstlike and of an all or nothing character. In the second group of interneurones the responses showed a quantitative dependence on the applied stimuli. In this group of interneurones responses often increased with the spatial extension of the peripheral stimulus revealing spatial summation of the afferent input.     

Effect of the mono- and tetra-sialogangliosides, GM1 and GQ1b, on long-term potentiation in the CA1 hippocampal neurons of the guinea pig

 Effects of the mono- and tetra-sialogangliosides, GM1 and GQ1b, on long-term potentiation (LTP) were investigated in the CA1 neurons of guinea-pig hippocampal slices. The magnitude of LTP induced by a strong tetanus (100 Hz, 100 pulses) was not significantly affected by application of either ganglioside. In contrast, when LTP was induced by a weak tetanus (100 Hz, 4 pulses), a significantly greater LTP was induced in the presence of either ganglioside. Similarly, when slices were incubated in low-Ca²⁺ (1.0–1.1 mM) medium for more than 2 h, the LTP was usually small or absent, but showed a significant increase in amplitude of population spike (A-PS) when the slices were incubated with either GM1 or GQ1b (4–5 μg/ml). In addition, the application of GQ1b (4 μg/ml) reversed the blocking effect of an NMDA-receptor antagonist, APP-5 (10 μM), on the induction of LTP and resulted in forming LTP. Based on these findings, we conclude that GM1 and GQ1b exert positive modulatory effects on the induction of LTP in hippocampal CA1 neurons and suggest that GM1 and GQ1b may participate in the induction of LTP as donors of Ca²⁺ ions. Received: 21 April 1998 / Accepted: 5 May 1998     

Hair-cell versus afferent adaptation in the semicircular canals

The time course and extent of adaptation in semicircular canal hair cells was compared to adaptation in primary afferent neurons for physiological stimuli in vivo to study the origins of the neural code transmitted to the brain. The oyster toadfish, Opsanus tau, was used as the experimental model. Afferent firing-rate adaptation followed a double-exponential time course in response to step cupula displacements. The dominant adaptation time constant varied considerably among afferent fibers and spanned six orders of magnitude for the population ( approximately 1 ms to >1,000 s). For sinusoidal stimuli (0.1-20 Hz), the rapidly adapting afferents exhibited a 90 degrees phase lead and frequency-dependent gain, whereas slowly adapting afferents exhibited a flat gain and no phase lead. Hair-cell voltage and current modulations were similar to the slowly adapting afferents and exhibited a relatively flat gain with very little phase lead over the physiological bandwidth and dynamic range tested. Semicircular canal microphonics also showed responses consistent with the slowly adapting subset of afferents and with hair cells. The relatively broad diversity of afferent adaptation time constants and frequency-dependent discharge modulations relative to hair-cell voltage implicate a subsequent site of adaptation that plays a major role in further shaping the temporal characteristics of semicircular canal afferent neural signals.     

Basic organization principles of the VOR: lessons from frogs

Locomotion is associated with a number of optical consequences that degrade visual information processing in the absence of appropriate compensatory movements. The resulting retinal image flow is counteracted by coordinated eye-head reflexes that are initiated by optokinetic and vestibular inputs. The contribution of the vestibulo-ocular reflex (VOR) for stabilizing retinal images is relatively small in amplitude in frogs but important in function by compensating for the non-linearities of the neck motor system. The spatial tuning of the VOR networks underlying the angular (AVOR) and linear (LVOR) with respect to canal and extraocular motor coordinates is organized in a common, canal-related reference frame. Thereby, the axes of head and eye rotation are aligned, principle and auxiliary VOR connections transform vestibular into motor signals and parallel AVOR and LVOR circuits mediate vergence and version signals separately. Comparison of these results with data from other vertebrates demonstrates a number of fundamental organization principles common to most vertebrates. However, the fewer degrees of behavioral freedom of frogs are reflected by the absence of, e.g. a functioning velocity storage network or of a fixation suppression of the VOR. In vitro experiments with the isolated brainstem and branches of N.VIII attached were used to study the putative transmitters of vestibular nerve afferent inputs, the postsynaptic receptor subtypes of second-order vestibular neurons and their dynamic response properties. Evidence is presented that suggests that afferent vestibular nerve fibers with different dynamic response properties activate different subtypes of glutamate receptors. The convergence pattern of monosynaptic afferent nerve inputs from different labyrinthine organs onto second-order vestibular neurons is remarkably specific. As a rule, second-order vestibular neurons receive converging afferent nerve inputs from one semicircular canal and from a specific sector of hair cells on one otolith organ. This convergence pattern remains malleable even in adulthood and reorganization is initiated by activity-related changes in vestibular nerve afferent fibers. The output of second-order vestibular neurons is modified by at least three inhibitory control loops. Uncrossed inhibitory vestibular side loops appear to control specifically the dynamic response tuning, whereas coplanar commissural inhibitory inputs improve mainly the spatial tuning and the cerebellar feedback loop controls the response gain. Among the targets of second-order vestibular projection neurons are extraocular motoneurons and internuclear neurons. Extraocular motoneurons differ among each other by the presence of very different response dynamics. These differences may represent a co-adaptation to the response dynamics of twitch and non-twitch extraocular muscle fibers. Different dynamical properties are required for a rapid acceleration of the globe at the one end and for the maintenance of a stable eccentric eye position over long periods of time at the other end of a continuum of variations in dynamic response properties. The maintenance of a given eccentric eye position over long periods of time is especially well developed in frogs and assists visual surveillance during lurking in the absence of saccades.     

Functional organization of the vestibular input to the anterior and posterior cerebellar vermis of cat

1. Responses evoked by electrical stimulation (auditory division of the VIIIth nerve sectioned chronically) and natural stimulation of the vestibular apparatus were recorded in the anterior and posterior cerebellar vermis of cats anesthetized with Ketamine or Nembutal. Under Ketamine the functional state of the cerebellar cortex was similar to that of the decerebrate or encéphale isolé preparation. 2. Vestibular-evoked responses were found bilaterally throughout the vermis (lob. I-X) and parts of pars intermedia and were, for the most part, mediated via the mossy fiber-granule cell pathway although natural stimulation occasionally evoked climbing fiber responses in Purkinje cells. 3. Lesion and stimulation experiments suggested that the polysynaptic potentials recorded in the dorsal folia of the anterior and parts of posterior vermis were mediated, at least in part, by the lateral reticular nucleus. Potentials recorded in the deeper folia often had shorter latencies and were probably mediated by primary and/or secondary vestibular fibers. Studies with horseradish peroxidase (injections in lob. V and VI) supported these notions. 4. An analysis of Purkinje cell responses to sinusoidal rotation and steps of angular acceleration or velocity indicated that P-cells in these regions signalled angular head velocity in the mid-frequency range. Single canal responses as well as multi-canal convergent P-cell responses were found. Purkinje cells also responded to static head displacement.     

Labyrinthine control of inferior oblique motoneurons

Summary Compensatory eye movements such as counterrolling are observed during head tilt. The role of each labyrinth and their bilateral subsequent interaction has been investigated by comparing the activity of motor nerve fibers innervating the inferior oblique (IO) muscle before and after hemilabyrinthectomy (HL). In encéphale isolé cats, the IO nerve was dissected and single or multiple unit activity was recorded. Discharge rate was computed and compared with the angular displacement of the head. The cat was placed on a rotating table which could rotate the head around an anterio-posterior (X) axis. Sinusoidal angular rotation (frequencies from 0.02 to 2 Hz, amplitude 0° to 30° peak to peak) was applied. Increase of unit activity was seen when the recorded side was moved upwards (SU). The maximum discharge rate lagged input velocity by 30° to 50°. Cessation of discharge occured during side down movement. Increase of activity during SU was attributed to excitation of the contralateral labyrinth (Vc) produced by activation of the vertical (mainly anterior) canals and utriculus. This excitation was augmented by the removal of inhibition arising from a decrease of activity in the vertical canals in the ipsilateral labyrinth (Vi).After acute HL on the Vi side a large increase of tonic background motoneuronal discharge was observed in the IO units. It may be due to a release from a) the direct inhibition which arises in Vi and b) commissural inhibition.The phasic character of the motor activity clearly present in the IL cats during tilt was not as sharp in the HL case.Its absence was attributed also to the lack of Vi inhibition which is assumed to reinforce the effect of excitation in the normal animals. Finally the nerve activity seems to lead velocity. This shift in the phase is similar to that previously observed in the neck and forelimb extensors. This data shows that the interaction between Vc and Vi plays an important role in the characteristics of the response, and suggests that inhibition contributes to the dynamic properties of the IO motoneurones compensatory action.A preliminary report of these results has been presented at the 2nd International Symposium on Motor Control. Varna. Bulgaria. October 1972.     

Reciprocal angular acceleration of the ankle and hip joints during quiet standing in humans

Human quiet standing is often modeled as a single inverted pendulum rotating around the ankle joint, under the assumption that movement around the hip joint is quite small. However, several recent studies have shown that movement around the hip joint can play a significant role in the efficient maintenance of the center of body mass (COM) above the support area. The aim of this study was to investigate how coordination between the hip and ankle joints is controlled during human quiet standing. Subjects stood quietly for 30 s with their eyes either opened (EO) or closed (EC), and we measured subtle angular displacements around the ankle (thetaa) and hip (thetah) joints using three highly sensitive CCD laser displacement sensors. Reliable data were obtained for both angular displacement and angular velocity (the first derivative of the angular displacement). Further, measurement error was not predominant, even among the angular acceleration data, which were obtained by taking the second derivative of the angular displacement. The angular displacement, velocity, and acceleration of the hip were found to be significantly greater (P<0.001) than those of the ankle, confirming that hip-joint motion cannot be ignored, even during quiet standing. We also found that a consistent reciprocal relationship exists between the angular accelerations of the hip and ankle joints, namely positive or negative angular acceleration of ankle joint is compensated for by oppositely directed angular acceleration of the hip joint. Principal component analysis revealed that this relationship can be expressed as: thetah=gammathetaa with gamma=-3.15+/-1.24 and gamma=-3.12+/-1.46 (mean +/-SD) for EO and EC, respectively, where theta is the angular acceleration. There was no significant difference in the values of y for EO and EC, and these values were in agreement with the theoretical value calculated assuming the acceleration of COM was zero. On the other hand, such a consistent relationship was never observed for angular displacement itself. These results suggest that the angular motions around the hip and ankle joints are not to keep the COM at a constant position, but rather to minimize acceleration of the COM.     

Receptive properties of sacral primary afferent neurons supplying the colon

1. Conscious perception of noxious and innocuous distension of the colon as well as the reflex control of anal continence and defecation largely depend on an intact sacral primary afferent innervation. Here we have studied the functional properties of these visceral primary afferent neurons in the dorsal root S2 in 17 cats. Single fibers projecting into the pelvic nerve were identified electrically and studied with innocuous and noxious mechanical stimulation of colon and anal canal. 2. A total of 59 units responding to one of these stimuli were investigated and they could be separated into two subpopulations of afferents. Thirty-six fibers were reproducibly excited by distension of the colon, but not by mechanical stimulation of the anal canal. They were thin myelinated or unmyelinated fibers with a median conduction velocity of 3.2 m/s. The remaining 23 units had receptive fields in the mucosa of the anal canal and responded readily to an innocuous proximodistal shearing stimulus, but not to distension stimuli applied to the same area. All, but two of these afferents were thin myelinated with a median conduction velocity of 7.7 m/s, which was significantly different from the conduction velocity of afferent neurons responding to distension of the colon. 3. Units responding to distension of the colon had thresholds in the innocuous range of the intracolonic pressure. Receptors that were activated only by noxious intraluminal pressure were absent. On the basis of their response to supramaximal isotonic distension, colonic afferents could be subclassified as phasic (n = 17) or tonic (n = 19) units. Phasic afferents were only transiently excited during filling or emptying of the colon, whereas tonic afferents discharged throughout the distension. The two populations had also significantly different median conduction velocities of 8.0 (n = 16) and 1.7 (n = 15) m/s, respectively. 4. Stimulation response functions were evaluated for 12 tonic afferents. All units encoded an increase of intracolonic pressure by the intensity of their discharge frequency. Increases of intracolonic pressure produced significantly higher discharge frequencies from unmyelinated than from thin myelinated afferents. 5. In three animals the percentage of unmyelinated fibers responding to mechanical stimulation of colon and anal canal was determined. Out of 213 electrically identified unmyelinated units projecting into the pelvic nerve, only 11 (5.2%) were excited. Thus, acute innocuous and noxious mechanical stimuli of the large intestine do not appear to be the adequate stimulus for the large majority of unmyelinated pelvic afferents. 6. In conclusion, distension of the colon and mechanical stimulation of the anal canal activates distinct populations of primary afferent neurons.(ABSTRACT TRUNCATED AT 400 WORDS)     

Asymmetric integration recorded from vestibular-only cells in response to position transients

Angular and translational accelerations excite the semicircular canals and otolith organs, respectively. While canal afferents approximately encode head angular velocity due to the biomechanical integration performed by the canals, otolith signals have been found to approximate head translational acceleration. Because central vestibular pathways require velocity and position signals for their operation, the question has been raised as to how the integration of the otolith signals is accomplished. We recorded responses from 62 vestibular-only neurons in the vestibular nucleus of two monkeys to position transients in the naso-occipital and interaural orientations and varying directions in between. Responses to the transients were directionally asymmetric; one direction elicited a response that approximated the integral of the acceleration of the stimulus. In the opposite direction, the cells simply encoded the acceleration of the motion. We present a model that suggests that a neural integrator is not needed. Instead a neuron with a long membrane time constant and an excitatory postsynaptic potential duration that increases with the firing rate of the presynaptic cell can emulate the observed behavior.     

Postnatal developmental changes in the response of rat primary horizontal semicircular canal neurons to sinusoidal angular accelerations

Summary At birth primary horizontal semicircular canal afferent neurons in the albino Wistar rat have slow, irregular spontaneous activity and insensitive, sluggish, variable responses to sinusoidal angular acceleration stimuli. There are rapid changes in the gross morphology of the rat semicircular canal in the first 4–5 days after birth, and during this time there is a rapid increase in neural gain re acceleration. Irregular neurons in rats about 6 days old have gains in the same range as irregular neurons in adult rats. However, after the gross morphological growth is complete, there continues to be a decrease in phase lag re acceleration. The causes of this developmental change in phase are unknown. It could be produced by changes in the receptor-afferent-efferent complex or by changes in the cupula or cupula-hair-cell attachment. These results with sinusoidal accelerations confirm the developmental increase in sensitivity and decrease in time constant found with constant angular accelerations (Curthoys 1979b).Supported by a grant from the National Health and Medical Research Council of Australia     

Variation in response dynamics of regular and irregular vestibular-nerve afferents during sinusoidal head rotations and currents in the chinchilla

In mammals, vestibular-nerve afferents that innervate only type I hair cells (calyx-only afferents) respond nearly in phase with head acceleration for high-frequency motion, whereas afferents that innervate both type I and type II (dimorphic) or only type II (bouton-only) hair cells respond more in phase with head velocity. Afferents that exhibit irregular background discharge rates have a larger phase lead re-head velocity than those that fire more regularly. The goal of this study was to investigate the cause of the variation in phase lead between regular and irregular afferents at high-frequency head rotations. Under the assumption that externally applied galvanic currents act directly on the nerve, we derived a transfer function describing the dynamics of a semicircular canal and its hair cells through comparison of responses to sinusoidally modulated head velocity and currents. Responses of all afferents were fit well with a transfer function with one zero (lead term). Best-fit lead terms describing responses to current for each group of afferents were similar to the lead term describing responses to head velocity for regular afferents (0.006 s + 1). This finding indicated that the pre-synaptic and synaptic inputs to regular afferents were likely to be pure velocity transducers. However, the variation in phase lead between regular and irregular afferents could not be explained solely by the ratio of type I to II hair cells (Baird et al 1988), suggesting that the variation was caused by a combination of pre- (type of hair cell) and post-synaptic properties.