Displacement configuration of semicircular canal cupulae.

Cupula displacement had traditionally been portrayed as a wiper motion of the cupula hinged at its base. In this study, semicircular canals of the bullfrog, which had been carefully dissected, were analyzed for cupula motion during compression of the canal wall. Cupulae marked with darkened oil droplets or alcian blue and displaced by endolymph exhibited a characteristic central bulging while the perimeter remained fixed to the ampullary wall. Small compressions resulted in displacements which were prominent near the receptor base while the center of the cupula mass had little movement. Large endolymph displacements produced a nearly symmetrical arching of the marker line that extended from the crista to the opposite wall. When endolymph was excessively displaced, the cupula became dislodged at its apical end and moved through the traditional wiper arc without the characteristic bulging.It is concluded that the cupula was dislodged in earlier studies by other investigators while, in the intact canal, the cupula has the properties of a modified elastic diaphragm. On the basis of these results, we suggest that the base of the cupula shears with the crista across the subcupular space.     

Effects of natural neck afferent stimulation on vestibulo-spinal neurons in the decerebrate cat

Summary The response patterns of antidromically identified vestibulo-spinal Deiters' neurons and other vestibular nuclear neurons to stimulation of neck receptors and horizontal semicircular canal receptors were investigated in precollicular decerebrate cats.The neck receptors were stimulated by sinusoidal oscillation (frequency 0.2 Hz and 10 ° amplitude) and trapezoidal movements of the cervical axis vertebrae in the horizontal plane, while the head was stereotaxically fixed in a prone position. The influence of horizontal semicircular canal receptors was examined by sinusoidal turntable oscillations (0.2 Hz, 10 °).Out of 151 antidromically driven Deiters' neurons, 16 units responded to neck rotation and 11 cells out of 148 Deiters' neurons were influenced by the horizontal semicircular canal receptors. Only 3 of these neurons were affected by convergent cervical and vestibular influences and revealed antagonistic response patterns. Neck-responsive units were found to be scattered throughout Deiters' nucleus, whereas semicircular-canal-influenced neurons were predominantly encountered in its rostro-ventral part.Altogether, 68 neck-responsive neurons in the vestibular nuclear complex were tested. These units showed a periodic, direction-specific modulation of the discharge rate in response to sinusoidal oscillation. Almost all responses were related to input angular velocity at 0.2 Hz with a mean gain of 1.15 imp/s per deg/s. During trapezoidal neck displacement, responsive units also exhibited tonical components in their discharge rates.These results suggest kinetic-static response characteristics of neck receptors. The cervico-vestibulo-spinal pathway may contribute to tonic neck reflexes; vestibular and neck reflexes may partially interact in Deiters' nucleus.Supported by Sonderforschungsbereich Hirnforschung und Sinnesphysiologie (SFB 70) der Deutschen Forschungsgemeinschaft     

Physiologic characteristics of vestibular first-order canal neurons in the cat. I. Response plane determination and resting discharge characteristics.

The response plane and resting rate characteristics of first-order, vestibular, semicircular canal neurons were studied in 67 cats under sodium pentobarbital anesthesia using single-unit recording techniques in the eighth nerve. Five hundred fifty-nine units were classified as to the canal they were associated with by employing an identification technique based on physiologic response patterns to brief, high-level (up to 250 degrees/S2) angular accelerations delivered in various head positions. All horizontal canal neurons had increased firing rates to ampullopetal and all vertical canal units to ampullofugal endolymph flow. The average observed roll and pitch null points for each canal were used to determine the average sensitivity vectors for the right horizontal, anterior, and posterior canals. These sensitivity vectors were at a variance of 4.6-10.2 degrees from those predicted by anatomical measurements (3). The mean resting discharge characteristics of 318 first-order neurons was 36.0 spikes/s (range 0.50-114 spikes/s). No significant difference was noted between horizontal and anterior canal neurons on horizontal and anterior canal neurons on the basis of resting rate. The resting rate of the posterior canal neuron population was significantly lower. The regularity of the resting discharge varied in all three canals and the average coefficient of variation was 0.238 for the population, with a range of 0.298-1.030. The population distribution of all resting-rate statistical parameters appeared to be unimodal, indicating that first-order canal neurons may not be broken into discrete populations on the basis of resting-rate characteristics. Of 47 adequately examined first-order canal neurons, 25 demonstrated a repeatable and predictable alteration in their resting discharge as their position to gravity was reoriented. This alteration was usually nonadapting and varied in magnitude according to the degree of tilt and original starting position. Of 25 canal gravity units, 4 had nearby units from the same canal which were unresponsive to gravity, suggesting the effect was due to a limited distortion of the crista or cupula rather than an overall displacement of the cupula.     

Determinants of spatial and temporal coding by semicircular canal afferents

The vestibular semicircular canals are internal sensors that signal the magnitude, direction, and temporal properties of angular head motion. Fluid mechanics within the 3-canal labyrinth code the direction of movement and integrate angular acceleration stimuli over time. Directional coding is accomplished by decomposition of complex angular accelerations into 3 biomechanical components-one component exciting each of the 3 ampullary organs and associated afferent nerve bundles separately. For low-frequency angular motion stimuli, fluid displacement within each canal is proportional to angular acceleration. At higher frequencies, above the lower corner frequency, real-time integration is accomplished by viscous forces arising from the movement of fluid within the slender lumen of each canal. This results in angular velocity sensitive fluid displacements. Reflecting this, a subset of afferent fibers indeed report angular acceleration to the brain for low frequencies of head movement and report angular velocity for higher frequencies. However, a substantial number of afferent fibers also report angular acceleration, or a signal between acceleration and velocity, even at frequencies where the endolymph displacement is known to follow angular head velocity. These non-velocity-sensitive afferent signals cannot be attributed to canal biomechanics alone. The responses of non-velocity-sensitive cells include a mathematical differentiation (first-order or fractional) imparted by hair-cell and/or afferent complexes. This mathematical differentiation from velocity to acceleration cannot be attributed to hair cell ionic currents, but occurs as a result of the dynamics of synaptic transmission between hair cells and their primary afferent fibers. The evidence for this conclusion is reviewed below.     

Response of semicircular canal dependent units in vestibular nuclei to rotation of a linear acceleration vector without angular acceleration

1. Recent experiments have shown that rotation of a linear acceleration vector round the head can generate involuntary ocular nystagmus in the absence of angular acceleration. The present experiments examine the suggestion that adequate stimulation of the semicircular canals may contribute to this response.2. Decerebrate cats were located in a stereotaxic device on a platform, slung from four parallel cables, which could be driven smoothly round a circular orbit without inducing significant angular movement of the platform. This Parallel Swing Rotation (PSR) generated a centripetal acceleration of 4.4 m/sec(2) which rotated round the head at 0.52 rev/sec.3. The discharge frequency of specifically lateral canal-dependent neural units in the vestibular nuclei of cats was recorded during PSR to right and left, and in the absence of motion. The dynamic responses to purely angular motion were also examined on a servo-driven turntable.4. Without exception all proven canal-dependent cells examined (twenty-nine cells in nine cats) were more active during PSR in the direction of endolymph circulation assessed to be excitatory to the unit, than during PSR in the opposite direction.5. The observed changes in discharge frequency are assessed to have been of a magnitude appropriate for the generation of the involuntary oculomotor response induced by the same stimulus in the intact animal.6. The findings suggest that a linear acceleration vector which rotates in the plane of the lateral semicircular canals can be an adequate stimulus to ampullary receptors, though an explanation which invokes the modulation of canal cells by a signal dependent upon the sequential activation of macular receptors cannot be positively excluded.     

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     

Interaction between the horizontal vestibulo-ocular reflex and optokinetic response in rabbits

Summary Dynamic characteristics of the horizontal vestibulo-ocular reflex (HVOR), the optokinetic response (OKR), and their interactions were investigated in alert albino rabbits. For stimulation of the horizontal semicircular canals, the whole rabbit was rotated sinusoidally on a motor-driven turntable at peak-to-peak amplitudes of 5 ° to 30 ° over a frequency range of 1/30 to 1/2 Hz. Optokinetic stimulation was provided by a narrow vertical slit light source presented in front of the eye to be tested. The evoked horizontal eye movements were observed and measured by means of a closed circuit television system adapted to provide an analog signal proportional to the eye movement. The net HVOR was obtained by rotation of the turntable in darkness and the net OKR by rotation of the light source. Combining rotation of the turntable with a stationary light source immediately increased the gain and reduced the phase shift of the HVOR. The light source moving in phase with the turntable, but at twice the angular amplitude, reduced the gain and advanced the phase of the HVOR. Eye movement curves of the HVOR modified by a fixed or moving slit light could be reconstructed approximately by a linear combination of the net HVOR and OKR.The work was supported by a grant from the Japanese Ministry of Education, Science and Culture (987008)On leave from C.N.R.S., France (from Nov. 1974 to Feb. 1975) and supported by a Franco-Japanese Cultural Relations Program FellowshipOn leave from C.N.R.S., France (from Nov. 1974 to Feb. 1975) and supported by a visiting professorship from Japan. Society for the Promotion of ScienceOn leave from Nencki Institute of Experimental Biology, Warsaw (from Jan. 1977 to Jan. 1978) and supported by a Polish-Japanese Cultural Relations Program Fellowship     

Some hydrodynamic properties of the posterior canal in the frog labyrinth related to neuronal responses

Measurements of the posterior-canal radius of curvature (R), the semicircular-duct radius (r) and the cupula radius (rC) were performed in the frog labyrinth. The Steinhausen-van Egmond equation led to an estimate of the canal endolymph flow, cupula deflection and sensory hair bending during constant angular accelerations (0.2-64 degrees/s2) of opposite directions and increasing duration (1.2-12 s). This analysis suggests that the non-linear canal afferent discharge behaviour exhibited by some units may not arise at the presynaptic level but rather postsynaptically, at the encoder site.     

Electrical activity of isolated semicircular canal in the frog

1. An isolated frog semicircular canal was placed in a two-compartment perspex chamber which allowed the fluids bathing the inside (endolymph) and the outside (perilymph) of the canal to be replaced separately. Mechanical stimulation (MS) of the ampullar receptors was performed by sinusoidal cupular movement.
2. Streptomycin sulfate in the perilymph reduced promptly, strongly and reversibly nerve spontaneous activity (SA) recorded on the ampullar nerve. Streptomycin sulfate also reduced reversible and weakly the slow potentials recorded either near the crista ampullaris (Adc) or on the ampullar nerve (Ndc) and the modulation of firing rate (ΔF_ms) all these activities being evoked by a mechanical stimulation.
3. Streptomycin sulfate injected into ampullar endolymph reduced promptly and strongly the Adc, Ndc, and ΔF_ms. When its concentration was higher 0.2 mg/ml, the SA was also reduced about 40 min later.
4. It is concluded that streptomycin sulfate alters weakly and reversibly spike generation process and synaptic transmission; it inhibits strongly and reversibly the resting discharge and it inhibits strongly and may be partly reversibly the transduction mechanisms.

     

人内耳半规管嵴顶时间常数的定量分析

目的 通过数值仿真和实验定量探究人内耳前庭半规管中的嵴顶时间常数,明确半规管编码角运动的时间过程。方法 建立人双耳半规管数值模型,通过流固耦合数值模拟嵴顶的生物力学响应,进而计算嵴顶的力学松弛时间常数。同时,对志愿者进行前庭眼反射实验,根据志愿者的眼震慢相角速度计算嵴顶的时间常数。结果 通过人内耳半规管数值模型计算得出的嵴顶力学松弛时间常数为3.75 s。通过实验测量得出平均嵴顶时间常数约为4.86 s。数值模型和实验中的结果近似保持一致。结论 人内耳前庭半规管中的嵴顶时间常数大约为4.86 s,反映了嵴顶力学松弛和半规管传入神经适应性的联合作用效果,体现了半规管编码角运动的时间过程。     

Principles of linear and angular vestibuloocular reflex organization in the frog.

We compared the spatial organization patterns of linear and angular vestibuloocular reflexes in frogs by recording the multiunit spike activity from cranial nerve branches innervating the lateral rectus, the inferior rectus, or the inferior obliquus eye muscles. Responses were evoked by linear horizontal and/or vertical accelerations on a sled or by angular accelerations about an earth-vertical axis on a turntable. Before each sinusoidal oscillation test in darkness, the static head position was systematically altered to determine those directions of horizontal linear acceleration and those planes of angular head oscillation that were associated with minimal response amplitudes. Inhibitory response components during angular accelerations were clearly present, whereas inhibitory response components during linear accelerations were absent. Likewise was no contribution from the vertical otolith organs (i.e., lagena and saccule) observed during vertical linear acceleration. Horizontal linear acceleration evoked responses that originated from eye muscle-specific sectors on the contralateral utricular macula. The sectors of the inferior obliquus and lateral rectus muscles on the utricle had an opening angle of 45 and 60 degrees, respectively and overlapped to a large extent in the laterorostral part of the utricle. Both sectors were coplanar with the horizontal semicircular canals. The sector of the inferior rectus muscle was narrow (opening 5 degrees), laterocaudally oriented, and slightly pitched up by 6 degrees. Angular acceleration evoked maximal responses in the inferior obliquus muscle nerve that originated from the ipsilateral horizontal and the contralateral anterior vertical canals in a ratio of 50:50. Lateral rectus excitation originated from the contralateral horizontal and anterior vertical semicircular canals in a ratio of 80:20. The excitatory responses of the inferior rectus muscle nerve originated exclusively from the contralateral posterior vertical canal. Measured data and known semicircular canal plane vectors were used to calculate the spatial orientation of maximum sensitivity vectors for the investigated eye muscle nerves in semicircular canal coordinates. Comparison of the directions of maximal sensitivity vectors of responses evoked by linear or angular accelerations in a given eye muscle nerve showed that the two vector directions were oriented about orthogonally with respect to each other. With this arrangement the linear and the angular vestibuloocular reflex can support each other dynamically whenever they are co-activated without a change in the spatial response characteristics. The mutual adaptation of angular and linear vestibuloocular reflexes as well as the differences in their organization described here for frogs may represent a basic feature common for vertebrates in general.     

Morphological correlates of response dynamics and efferent stimulation in horizontal semicircular canal afferents of the toadfish, Opsanus tau.

1. We used the intraaxonal labeling technique to study correlations between the terminal dendritic morphology of horizontal semicircular canal primary afferents and their response dynamics to sinusoidal head rotation and combined electrical stimulation of central efferent vestibular neurons. Thirty-eight canal afferents were identified by their sensitivity and phase of response to rotation between 0.1 and 1.0 Hz (+/- 10 degrees/s) and were subsequently labeled with horseradish peroxidase or biocytin. The afferent's dendritic field and synaptic specializations in the neuroepithelium of the crista were examined under light microscopy. 2. Rate and regularity of background discharge of the afferent were not correlated with its axon diameter or relative location of its dendritic field in the crista. 3. Response sensitivity of the afferent to rotation was correlated both with the relative location of its dendritic field in the crista and with the number of terminal endings it possesses. Afferents having low sensitivities, slow dynamics, and few terminal endings supply the peripheral portions of the crista; afferents with higher sensitivities, faster dynamics, and greater number of terminal endings supply the more central portions. It is suggested that the differences in sensitivity among the afferents reflect principally the variations in both the cupular dynamics along the crista and the number of possible hair cell contact sites in the neuroepithelium. 4. Response phase of the afferent was correlated only with the extent of its dendritic processes along the transverse axis of the crista. Afferents having transversely oriented dendritic fields had less phase lags relative to acceleration than did those having a more longitudinally oriented dendritic field. 5. Efferent stimulation produced a change in both the afferent's discharge rate and its response sensitivity to rotation. Afferents discharge rate and its response sensitivity to rotation. Afferents having a centrally located dendritic field and acceleration afferents, defined by their response to rotation, were the most affected by efferent stimulation. These results suggest that efferent innervation is either directed toward, or most efficacious in, the central regions of the crista and that it may select specific hair cell-afferent complexes.     

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.     

上半规管良性阵发性位置性眩晕诊断试验分析和物理引擎耳石运动观察

目的 对上半规管良性阵发性位置性眩晕（BPPV）诊断试验进行分析，了解其诊断机制。方法 建立标准空间坐标系的膜迷路模型，设定膜迷路不同位置结石，基于物理引擎三维物理仿真，分析Dix Hallpike试验和仰卧悬挂头位试验耳石运动情况，进而推断所诱发眼震表现。结果 Dix-Hallpike试验和仰卧悬挂头位试验所有位置的结石都有一定程度的运动，并能诱发椭圆囊的结石经总管进入半规管。出现下跳眼震的情况，包括结石从椭圆囊进入后半规管，对侧后半规管结石向壶腹运动，上半规管的结石从壶腹嵴帽底部滑动到顶部，后者重复诊断实验仍为下跳眼震。重复10次观察，试验结果一致。结论 对于上半规管BPPV的诊断标准，还需要重新评估，观察眼震动态变化是必要的，其诊断方法需要改良。     

外半规管良性阵发性位置性眩晕诊断试验分析和物理引擎耳石运动观察

目的对外半规管良性阵发性位置性眩晕(BPPV)诊断试验进行分析,了解其诊断机制。方法建立基于物理引擎三维物理仿真的BPPV迷路模型,设定外半规管不同位置结石,分析水平滚转试验过程不同位置耳石运动情况,进而可以推断所诱发眼震表现。结果水平滚转试验眼震表现形式多样,包括双侧背地眼震,可以判断为壶腹部结石和嵴帽结石,眼震弱侧为患侧;双侧向地眼震,可以判断为外半规管长臂侧结石,眼震强烈侧为患侧;一侧背地眼震一侧向地眼震,考虑长臂侧壶腹部结石;水平滚转试验存在的设计缺陷是会导致耳石复位,影响诊断试验的敏感性。10次重复试验结果一致。结论60°水平滚转试验有效修正了90°水平滚转试验存在的缺陷。基于物理引擎对诊断试验进行分析,对于我们理解外半规管BPPV的诊断机制有重要意义,也有助于改良和创新诊断方法。     

Responses of pigeon horizontal semicircular canal afferent fibers. I. Step, trapezoid, and low-frequency sinusoid mechanical and rotational stimulation

1. The horizontal semicircular canals of anesthetized (barbiturate/ketamine) pigeons were stimulated by rotational and by mechanical stimulation. 2. The mechanical stimulation consisted of making a small (less than 1 mm) fistula in the lateral part of the bony horizontal semicircular canal and, after inserting a probe coupled to a piezoelectric micropusher through the fistula, providing controlled indentation of the exposed membranous horizontal semicircular duct. 3. Extracellular action potentials from single horizontal semicircular canal primary afferent (HCA) fibers were recorded during sinusoidal rotational and during step, ramp, and sinusoidal mechanical stimulation. 4. The mean spontaneous discharge rate of 160 horizontal canal afferents was 86 +/- 4 (SE) spikes/s. This rate was not significantly different from that reported previously for pigeon HCA fibers recorded with the horizontal canal intact (i.e., no fistula introduced). 5. Sinusoidal mechanical indentation of the horizontal semicircular duct produced clearly entrained action potentials on 36 HCA fibers for a range of peak displacements from +/- 0.5 to +/- 30 microns. Action potentials were never modulated on afferents (n greater than 100) identified as innervating the anterior and posterior semicircular canals or the otolith organs during mechanical stimulation of the horizontal semicircular canal, even for displacements as large as 30 microns. 6. Intensity functions relating peak firing frequency (spikes per second) and peak probe displacement (micrometers) for 1.0-Hz sinusoidal mechanical stimulation were linear over the range 1.0-5.0 microns. The most sensitive units (6/36, 17%) showed response saturation as the stimulus magnitude was extended to 7 microns and beyond. 7. In 15 of 36 units, both mechanical and rotational sinusoidal stimulation (1.0 Hz) were applied to the same unit. The duct indentation magnitudes were 1.0, 2.5, 5.0, and 7.0 microns and the rotational velocities were 5, 10, and 20 deg/s. The constant of proportionality found to equate the peak response produced by rotational to that elicited by mechanical stimulation was 7.0 deg.sec-1/1.0 microns. 8. Bode plots and best-fit transfer functions of the frequency response (0.05-10.0 Hz) of 14 HCAs exposed to both mechanical and rotational stimulation were nearly identical. 9. Parameters for best-fit transfer functions, responses to step, and trapezoidal duct displacements were in excellent agreement with previous rotational studies carried out using the pigeon. 10. Although the mechanisms by which focal identation of the horizontal membranous duct produce responses have not yet been determined, primary afferent responses using this method of stimulation are directly comparable with rotatory stimulation.(ABSTRACT TRUNCATED AT 400 WORDS)     

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.     

Unilateral testing of utricular function

A modified rotatory chair test is reported in which radial acceleration, generated by eccentric displacement of the subject during constant angular velocity, is exploited as a unilateral stimulation to the otolith organs. During constant angular rate rotation, the test subject is displaced laterally on the rotating turntable by 3.5 cm, so that one labyrinth becomes aligned with the rotatory axis while the second – eccentric – labyrinth is solely exposed to the altered gravito-inertial acceleration (GIA). Previously reported results showed that the direction of the response is independent of the direction of turntable rotation, ruling out any canal influence, and indicated that in a normal population the response, measured in one eye, was symmetrical for displacement of the left and right labyrinths. This mode of stimulus thus appears to elicit a unilateral otolith-ocular response (OOR). Examination of this unilateral OOR was extended in the present study; comparative testing with head-tilt to gravity, i.e. involving bilateral stimulation to the otolith organs, was carried out. Movements of both eyes were recorded (by three-dimensional video-oculography), in order to examine response conjugacy. To verify the specificity of the unilateral stimulus, tests were performed with patients who had previously undergone unilateral section of the vestibular nerve as treatment for acoustic neuroma. The eccentric displacement profile (EDP) and head-tilt stimulus each included ten cycles of left-right oscillation in order to permit signal averaging. In the normal subjects (n=12) the torsional component of the OOR proved to be both labyrinth-symmetrical and conjugate, during both bilateral and unilateral otolith stimulation. OOR gain (ocular torsion/GIA tilt) was higher for bilateral than unilateral stimulation. Bilateral OORs, obtained from three of the five unilaterally deafferented patients, proved less symmetrical and conjugate than in the normals. Unilateral OORs in all five patients were characteristically asymmetrical, with little or no response during stimulation of the diseased labyrinth. Received: 28 July 1997 / Accepted: 3 February 1998     

Cupula displacement,hair bundle deflection,and physiological responses in the transparent semicircular canal of young eel

The transparent labyrinth of young eels (Anguilla anguilla L.) was used in toto for studying the configuration of cupula displacement, deflection of the hair bundle, and correlated changes in transepithelial voltage (TEV) and nerve activity (NA) in the semicircular canal. Microcapillaries were introduced into the canal through holes produced by a microthermocauter. Mechanical stimulation was applied either by injection of fluid into the ampulla or by electromagnetically displacing ferrofluid as a piston within the canal. Motion of individual kinocilia, stained cupulae or the ferrofluid piston was analysed by double-exposed microphotographs, photodiodes, or a video-system. The three-dimensional cupula displacement configuration was found to be piston-to diaphragm-like. Hair bundles at different sites on the crista exhibit differences in amplitude and time course of deflection. The transfer factor between shifts of the canal fluid and the tips of the kinocilia is 0.4–0.6. Displacements in opposite directions induce TEV and NA of opposite sign. Various tests confirmed TEV to reflect receptor potential responses. Nerve activity adapts to a tonic response with a time constant of 6.4s. No similar adaptation occurred in TEV. Stimulus-response curves of TEV- and NA-responses are similar and signoid in shape with saturation at ciliary deflections of roughly + 6° and –3°.