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Direct and indirect pathways mediating the vestibulo-ocular reflexes, considering contributions made by the vestibular commissure, cerebellum, and vestibular efferent neurons are reviewed. This background leads to a detailed treatment of three-dimensional aspects of the vestibulo-ocular reflex, comparing the planes of the labyrinthine semicircular canals with the pulling directions of the extraocular muscles. By applying matrix algebra and rotational vectors, Robinson provides insights into the comparative anatomy of the vestibular system in different species, how central circuits process raw vestibular signals in three dimensions, and how the directions of eye movement caused by brainstem and cerebellar lesions can be explained., (Copyright © 2022 Elsevier B.V. All rights reserved.)

The biophysical properties of the labyrinthine semicircular canals, and the electrophysiological properties of peripheral vestibular afferent neurons over a range of stimulus frequencies, are reviewed. Resting discharge activity and adaptive properties of vestibular neurons are discussed. Central processing of vestibular signals is then examined, including push-pull organization and the velocity storage mechanism. A detailed treatment of the final common neural integrator for oculomotor signals follows with consideration of its neural substrate and how distributed networks of neurons can overcome several problems posed by conventional control-systems models, such as why neural signals, but not background discharge, are integrated. Next, the behavior of the vestibulo-ocular reflex in darkness is compared with how it satisfies visual demands during natural activities. Finally, the reflex's performance at high frequencies of head rotation is discussed., (Copyright © 2022 Elsevier B.V. All rights reserved.)

Aims: Although the vestibular system has been widely investigated over the past 50 years, there is still an unsolved mystery. Some special vestibular afferent (SVA) neurons responding to both rotation and linear force were found through neurophysiological techniques, however, the sensory overlap mechanism of SVA neurons is still unclear, which may be closely related to vestibular-related diseases., Materials and Methods: To address the above-mentioned problem, a cupula buoyancy theory was established in the present study, where SVA neurons were considered semicircular canal afferent (SCCA) neurons. Then labyrinth anatomy and neural response dynamics of vestibular afferent neurons in chinchilla were investigated through vestibular labyrinth reconstruction and single unit recording technique, respectively., Key Findings: We analyzed the deflections of cupulae under multiple conditions with the help of Amira Software and predicted the neural response law of SCCA neurons to linear force based on the cupula buoyancy theory. Data analysis confirmed that the basic response characteristic of SVA neurons had no significant difference to those of SCCA neurons, but were significantly different from those of otolith afferent neurons. Further, the actual responses of SVA neurons to linear force are completely consistent with our predictions. These results strongly suggest that SVA neurons actually are SCCA neurons, and the cupula buoyancy theory is the key to the sensory overlap mechanism of SCCA neurons., Significance: Our study revealed the real identity of SVA neurons and provided a reasonable mechanism for sensory overlap of rotation and linear force, which improved our understanding about the vestibular system., Competing Interests: Declaration of competing interest The authors declare that they have no competing interests, and all authors confirmed the accuracy of the data., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Objective: The present study was designed to determine whether healthy older adults with age-related vestibular loss have deficits in spatial navigation., Methods: 154 adults participating in the Baltimore Longitudinal Study of Aging were tested for semicircular canal, saccular, and utricular function and spatial navigation ability using the blindfolded Triangle Completion Test (TCT). Multiple linear regression was used to investigate the relationships between each measure of vestibular function and performance on the TCT (angular error, end point error, and distance walked) while controlling for age and sex., Results: Individuals with abnormal saccular function made larger angular errors (β = 4.2°, p < 0.05) and larger end point errors (β = 13.6 cm, p < 0.05). Independent of vestibular function, older age was associated with larger angular (β's = 2.2-2.8°, p's < 0.005) and end point errors (β's = 7.5-9.0 cm, p's < 0.005) for each decade increment in age., Conclusions: Saccular function appears to play a prominent role in accurate spatial navigation during a blindfolded navigation task., Significance: We hypothesize that gravitational cues detected by the saccule may be integrated into estimation of place as well as heading direction., (Copyright © 2019 International Federation of Clinical Neurophysiology. All rights reserved.)

Commissural connections between the superior colliculi (SCs) were well known anatomically, and assumed to be only inhibitory in relation to visual inputs. However, by recording intracellular potentials, we revealed that a strong monosynaptic excitatory commissural connection exists between the rostral SCs of the cat. Commissural excitation existed between the medial-medial or lateral-lateral parts of both SCs, while commissural inhibition existed between the medial SC on one side and the lateral SC on the opposite side. These commissural excitation and inhibition were also confirmed morphologically with the double-labeling method of HRP-conjugated gold particle and GABA. Similarity of the topography of commissural inhibition between the SC system and the vestibuloocular system supported the conclusion that the saccadic eye movement system uses the same semicircular canal coordinate as the vestibuloocular system. The commissural excitation may help to maintain Listing's law in saccadic eye movements., (© 2019 Elsevier B.V. All rights reserved.)

Objective: We aimed to quantify peripheral-vestibular deficits that may contribute to imbalanced stance/gait in patients with inflammatory neuropathies., Methods: Twenty-one patients (58 ± 15 y [mean age ± 1SD]; chronic-inflammatory-demyelinating-polyneuropathy = 10, Guillain-Barré Syndrome = 5, Anti-MAG peripheral neuropathy = 2, multifocal-motor-neuropathy = 4) were compared with 26 healthy controls. All subjects received video-head-impulse testing (vHIT), caloric irrigation and cervical/ocular vestibular-evoked myogenic-potentials (VEMPs). The Yardley vertigo-symptom-scale (VSS) was used to rate vertigo/dizziness. Postural stability was assessed using the functional gait-assessment (FGA). Pure-tone audiograms (n = 18), otoacoustic emissions (n = 12) and auditory brainstem responses were obtained (n = 12)., Results: Semicircular-canal hypofunction was noted in 9/21 (43%) patients (vHIT = 6; caloric irrigation = 5), whereas otolith function was impaired in 12/21 (57%) (oVEMPs = 8; cVEMPs = 5), resulting in vestibular impairment of at least one sensor in 13/21 (62%). On average, 2.4 ± 1.1 vestibular end organs (each side: anterior/posterior/horizontal canal, utriculus, sacculus; total = 10) were affected. The VSS-scores were higher in patients (16.8 ± 8.6 vs. 9.5 ± 6.2, p = 0.002) but did not correlate with the number of affected organs. Auditory neuropathy was found in 1/12 (8%) patients., Conclusion: Impairment of one or more vestibular end organs was frequent, but usually mild, possibly contributing to imbalance of stance/gait in inflammatory neuropathies., Significance: While our data does not support routine vestibular testing in inflammatory neuropathies, this may be considered in selected cases., (Copyright © 2017 International Federation of Clinical Neurophysiology. Published by Elsevier B.V. All rights reserved.)

Objective: To explore ocular vestibular evoked myogenic potentials (oVEMP) to low-frequency vertex vibration (125 Hz) as a diagnostic test for superior canal dehiscence (SCD) syndrome., Methods: The oVEMP using 125 Hz single cycle bone-conducted vertex vibration were tested in 15 patients with unilateral superior canal dehiscence (SCD) syndrome, 15 healthy controls and in 20 patients with unilateral vestibular loss due to vestibular neuritis. Amplitude, amplitude asymmetry ratio, latency and interaural latency difference were parameters of interest., Results: The oVEMP amplitude was significantly larger in SCD patients when affected sides (53 μVolts) were compared to non-affected (17.2 μVolts) or compared to healthy controls (13.6 μVolts). Amplitude larger than 33.8 μVolts separates effectively the SCD ears from the healthy ones with sensitivity of 87% and specificity of 93%. The other three parameters showed an overlap between affected SCD ears and non-affected as well as between SCD ears and those in the two control groups., Conclusions: oVEMP amplitude distinguishes SCD ears from healthy ones using low-frequency vibration stimuli at vertex., Significance: Amplitude analysis of oVEMP evoked by low-frequency vertex bone vibration stimulation is an additional indicator of SCD syndrome and might serve for diagnosing SCD patients with coexistent conductive middle ear problems., (Copyright © 2016 International Federation of Clinical Neurophysiology. Published by Elsevier Ireland Ltd. All rights reserved.)

In the last two decades, we have focused on establishing a reliable technique for focal stimulation of vestibular receptors to evaluate neural connectivity. Here, we summarize the vestibular-related neuronal circuits for the vestibulo-ocular reflex, vestibulocollic reflex, and vestibulospinal reflex arcs. The focal stimulating technique also uncovered some hidden neural mechanisms. In the otolith system, we identified two hidden neural mechanisms that enhance otolith receptor sensitivity. The first is commissural inhibition, which boosts sensitivity by incorporating inputs from bilateral otolith receptors, the existence of which was in contradiction to the classical understanding of the otolith system but was observed in the utricular system. The second mechanism, cross-striolar inhibition, intensifies the sensitivity of inputs from both sides of receptive cells across the striola in a single otolith sensor. This was an entirely novel finding and is typically observed in the saccular system. We discuss the possible functional meaning of commissural and cross-striolar inhibition. Finally, our focal stimulating technique was applied to elucidate the different constructions of axonal projections from each vestibular receptor to the spinal cord. We also discuss the possible function of the unique neural connectivity observed in each vestibular receptor system., (Copyright © 2011 Elsevier Ireland Ltd and the Japan Neuroscience Society. All rights reserved.)

After acute vestibular loss in humans or animals, eye-movement responses to rapid horizontal ipsilesional head rotations ("head impulses") show that there is severe, permanent impairment of the angular vestibulo-ocular reflex. The basis for this appears to be an inhibitory saturation of ipsilesional vestibular nerve, and perhaps vestibular nucleus afferents, in response to high-acceleration, off-direction stimulation of the sole functioning member of any pair of semicircular canals. For the clinician, impulsive testing is an easy and reliable way to identify severe unilateral as well as bilateral impairment of semicircular canal function.

Convergent inputs from the lateral and vertical semicircular canals (LC and VC) to 31 central vestibular-only (VO) and vestibular-plus-saccade (VPS) neurons were determined by oscillating monkeys about a spatial vertical axis while the head was tilted forward and backward up to 90 degrees. Activity of each neuron varied as a function of head tilt. Seven neurons had maximal activation when the head was tilted approximately 30 degrees forward (spatial phase), indicating convergent inputs from the LC, while peak activation of 10 units occurred with the head tilted back approximately 50 degrees, indicating VC input. Fourteen neurons had spatial phases that deviated more that 15 degrees from the LC and VC planes, indicating convergent inputs from LC and VC. Seven of these neurons had a spatial phase less than 15 degrees forward and 35 degrees back, indicating canal inputs from both sides. Seven other neurons had spatial phases more that 45 degrees forward and 65 degrees back, indicating inputs from canals located on the same side. Thus, there are two groups of central vestibular neurons: one group responds maximally when the head is rotated about a spatial vertical axis in an upright position, declining as the head is tilted away from this position. Another group responds minimally to rotation in an upright head orientation, increasing as the head is tilted away from the upright. A majority of the cells also had convergent otolith input. The otolith and canal inputs superposed when the animals were rotated about roll and pitch axes from an upright position. This insured that these neurons would respond over a broad frequency range from very low to high frequencies.

During constant-velocity rotation about a tilted axis (OVAR), the VOR and the rotation perception last indefinitely, but show a striking dependency on tilt angle. We show that, during OVAR, a variety of motions can account for the head motion relative to gravity. Some of these are in conflict with canal signals, but correspond to a lower angular velocity; we suggest that the brain performs a trade-off in order to select the best motion. We show that this theory explains the effect of tilt angle on velocity estimation during OVAR.

Three subnuclei within the inferior olive are implicated in the control of eye movement; the dorsal cap (DC), the beta-nucleus and the dorsomedial cell column (DMCC). Each of these subnuclei can be further divided into clusters of cells that encode specific parameters of optokinetic and vestibular stimulation. DC neurons respond to optokinetic stimulation in one of three planes, corresponding to the anatomical planes of the semicircular canals. Neurons in the beta-nucleus and DMCC respond to vestibular stimulation in the planes of the vertical semicircular canals and otoliths. Each these olivary nuclei receives excitatory and inhibitory signals from pre-olivary structures. The DC receives excitatory signals from the ipsilateral nucleus of the optic tract (NOT) and inhibitory signals from the contralateral nucleus prepositus hypoglossi (NPH). The beta-nucleus and DMCC receive inhibitory signals from the ipsilateral nucleus parasolitarius (Psol) and excitatory signals from the contralateral dorsal Y group. Consequently, the olivary projection to the cerebellum, although totally crossed, still represents bilateral sensory stimulation. Inputs to the inferior olive from the NOT, NPH, Psol or Y-group discharge at frequencies of 10-100 imp/s. CFRs discharge at 1-5 imp/s; a frequency reduction of an order of magnitude. Inferior olivary projections to the contralateral cerebellum are sagittally arrayed onto multiple cerebellar folia. These arrays establish coordinate systems in the flocculus and nodulus, representing head-body movement. These climbing fiber-defined spatial coordinate systems align Purkinje cell discharge onto subjacent cerebellar and vestibular nuclei. In the oculomotor system, olivo-cerebellar circuitry enhances and modifies eye movements based on movement of the head-body in space.

Objective: To determine the effects of stimulation of bilateral lateral semicircular canals (LSCCs) by accelerated rotation and caloric stimulation of unilateral LSCC on vestibular evoked myogenic potentials (VEMPs) in healthy volunteers., Methods: In experiment 1, VEMPs were recorded while subjects (n = 11) were seated in a rotational chair and angular acceleration around the earth-vertical axis was provided. Amplitudes of p13-n23 were corrected using background muscle activities. In experiment 2, subjects (n = 8) in the semilateral position kept the LSCC in the vertical position and activated the sternocleidomastoid muscle by twisting the neck. After irrigating the external auditory canal with ice water, VEMPs were recorded on the irrigated side. In experiment 3, the same setting as experiment 2 was applied (n = 6), and hot water of 44 degrees C was used for irrigation., Results: There were no significant differences in latencies of p13 or n23, and in corrected amplitudes by either rotatory or caloric stimulation., Conclusions: Simultaneous stimulation of LSCCs has little effect on VEMPs., Significance: No functional interaction between the saccule and LSCC was detected in VEMPs, although convergence of semicircular canal and otolith nerve inputs onto single vestibular nucleus neurons has been demonstrated electrophysiologically in animal experiments.

In this chapter, we describe our recent work on the divergent properties of single, long descending motor-tract neurons in the spinal cord, using the method of intra-axonal staining with horseradish peroxidase, and serial-section, three-dimensional reconstruction of their axonal trajectories. This work provides evidence that single motor-tract neurons are implicated in the neural implementation of functional synergies for head movements. Our results further show that single medial vestibulospinal tract (MVST) neurons innervate a functional set of multiple neck muscles, and thereby implement a canal-dependent, head-movement synergy. Additionally, both single MVST and reticulospinal axons may have similar innervation patterns for neck muscles, and thereby control the same functional sets of neck muscles. In order to stabilize redundant control systems in which many muscles generate force across several joints, the CNS routinely uses a combination of a control hierarchy and sensory feedback. In addition, in the head-movement system, the elaboration of functional synergies among neck muscles is another strategy, because it helps to decrease the degrees of freedom in this particularly complicated control system.

The endolymph flow inside the semicircular ducts is analytically investigated by considering a system of two hydrodynamically interconnected ducts. Rotation of this system adds an amount of motion (momentum) to parts of it. This results in an endolymph flow in generally all vestibular parts. The "external impulses" are the impulses which emerge by rotation of exclusively a particular vestibular part. The real impulses can be calculated from a set of equations which contain the external impulses. Analytical expressions are derived for the initial velocities in the ducts and for the maximum endolymph displacements. These formulae contain the external impulses and the ratios of: (1) the radii of crus commune and ducts (gamma), (2) the lengths of crus commune and ducts (lambda). It was proven that an interconnected system composed of two ducts, and also a system composed of two such semicircular duct systems, behaves as a pure rotation transducer (like a single duct does), also when it is rotated excentrically. Duct systems with polygonal and circular geometries were used to evaluate whether an optimal value of lambda would exist (gamma was already considered elsewhere). Optimum values of lambda in a range of about 0.10-0.52 were found. This rather wide range of values agrees with values from measurements. Optimization of an interconnected duct system appeared to be equal to optimization of a system composed of separate ducts.

In the majority of vertebrates, the horizontal duct of the vestibular system lies approximately in the yawing plane of the head. The positioning of the vertical ducts, however, is not in the pitch- and roll planes but the vertical ducts generally lie under an angle of about 30-45 degrees relative to the medial plane. Using the equations for a hydrodynamically interconnected two-duct system, optimal positions of the vertical and horizontal ducts in different vertebrate groups can be derived. It was stated that the mean response of the vertical ducts should be optimized. This leads to a symmetrical positioning of the vertical ducts with respect to the medial plane. In all observed vertebrate groups, a solution of mu =(pi-alpha)/2 is found (mu is the angle of the vertical ducts relative to the medial plane, alpha is the angle between the vertical duct planes). For alpha=90 degrees, this provides an equal sensitivity for pitch- and roll- movements. For alpha>90 degrees, a larger sensitivity for pitch movements is obtained, at the expense of a lower sensitivity for roll movements. It is argued that the angle alpha between the vertical ducts may vary from 90 to 120 degrees. In most vertebrates, the centre of mass is stabilized by e.g. fins, tri- or quadrupedal stability, a crawling body or upside-down resting positions (e.g. bats). Birds are generally biped, so in walking they are also rather sensitive to roll. These features are related to labyrinth positioning in the head.

The classical representation of the semicircular duct system consists of three separate duct circuits. The ducts are, however, in reality, hydrodynamically interconnected. Muller & Verhagen (1988a,b) derived equations for the mechanical behaviour of an interconnected system with three ducts (anterior, posterior and horizontal). An analytical solution of these equations would, however, be too complex to provide surveyable formulae. A system of two interconnected ducts avoids this complexity whilst keeping the essentials of the coupling of ducts intact. The solution of the equation of motion leads to expressions for time constants and maximal endolymph excursions which are functions of morphological parameters, viz. the ratios of radii (gamma) and lengths (lambda) of the common vestibular part (crus commune or utriculus) and the ducts. The system possesses two short time constants which are shown to have similar values. The maximum endolymph displacements in the two ducts after a steplike stimulus are the products of the respective initial velocities and combinations of time constants. The initial velocities depend strongly on the position of the labyrinth with respect to the excitating rotation vector. Measured data of gamma and lambda are compared with the theoretical results. For gamma, excellent agreement was found. lambda is treated elsewhere.

Frogs and toads turn either their heads or bodies opposite to angular accelerations applied around the yaw axis. Thresholds exist for the minimum angular acceleration that induces this vestibulomotor response in individual frogs. These thresholds were recorded for several anuran species that cover a broad range of sizes and life styles. Interspecific variation in the magnitude of the thresholds, which correlated with the ecology and behavior of the species, was documented. Also an allometric relationship was observed between this threshold and body size; the larger the frog, the lower the threshold. In many species, the threshold value for reflexive vestibulomotor responses to angular acceleration was proportional to the -0.4 (+/-0.2) power of body mass. Physical dimensions of the semicircular canals determine, in part, vestibular sensitivity to angular acceleration. Hence changes with growth in the semicircular canals are believed to contribute to the slope of -0.4. The biological significance of this allometry in vestibular responses is discussed and compared to trends in vestibular sensitivity and semicircular canal morphology of other vertebrate classes., (c1999 COSPAR. Published by Elsevier Science Ltd.)

Objective: The aim of this study was to determine the otolithic and semicircular canal effects of galvanic vestibular stimulation with increasing current strengths on eye movements and the perception of verticality., Methods: We measured (1) 3-dimensional eye movements, (2) subjective tilt of the peripheral visual field, and (3) subjective tilt of a central vertical line in 12 healthy subjects during galvanic vestibular stimulation. A rectangular, unipolar binaural electric current was applied to each subject's mastoid., Results: Anodal stimulation of the right mastoid led to an ipsiversive tonic ocular torsion of up to 5.4 degrees, to a contralateral tilt of both the peripheral visual field (1-9 degrees), and a central vertical line (0.5-6.2 degrees) increasing in amplitude with increasing current strengths applied. This reflects otolith stimulation. In most subjects, current strengths of 3 mA or more elicited a slight (horizontal-) torsional nystagmus (amplitude 1-2 degrees) that was superimposed on static torsion. This reflects horizontal and vertical semicircular canal stimulation. A correlation was found in the amount of the 3 measured parameters and the strength of the applied current., Conclusions: Thus, galvanic vestibular stimulation at low current intensities (1-3 mA) preferably excites otolith responses, which increase with increasing current intensity. With higher current intensity above 3 mA, additional semicircular canal responses are elicited in the form of horizontal-rotatory nystagmus superimposed on static torsional deviations. The lack of a vertical deviation and nystagmus can be explained by the counterdirected vertical components of the anterior and posterior semicircular canal.

Pre-mated adult female newts and fertilized eggs will be flown on the International Microgravity Laboratory-2 flight, in 1994. One objective of the flight will be to observe the influence of microgravity on the development of the gravity-sensing organs in the inner ear. These organs contain sensory hair cells covered by a layer of dense stones (otoconia). Gravity and linear acceleration exert forces on these masses, leading to excitation of the nerve fibers innervating the hair cells. If the production of the otoliths is regulated to reach an optimal weight, their development might be abnormal in microgravity. Ground-based control experiments are reported describing the developmental sequence in which both the otoliths and their associated sensory epithelium and the semicircular canals appear and develop. Three-dimensional reconstruction of serial sections through the otic vesicle of newt embryos at stages 31 through 58 demonstrate the first appearance, relative position and growth of the otoliths. Reports of experiments in which fertilized frog eggs were flown on a Russian Cosmos mission conclude that the utricular otolith is increased in volume, whereas the saccular otolith maintains normal size, suggesting that at least in the utricle, the weight of the otolith might be regulated.

It gets into difficult to explain the mechanism of caloric nystagmus only by convection theory from results of microgravity experiments. One of the other theories is an occurrence of a relative volume change due to a temperature change. Since the volume change must lead to a pressure change after caloric stimulation, we tried to measure the ampulla pressure of the horizontal semicircular canal in pigeons (Columba livia) using an improved servo micropipette system. The main result was that the ampulla pressure increased by cooling and decreased by heating. The changes of the ampulla pressure depended on the temperature change but were not influenced by the pigeon's head position.

Short latency vestibular evoked potentials (VsEPs) to angular acceleration impulses (maximal intensity 20,000 degrees/sec2, rise time 1.5-3 msec) were recorded by skin electrodes in cats before and after various surgical procedures. Under general anesthesia, the animals underwent unilateral labyrinthectomy and the VsEPs in response to stimulation of the remaining inner ear in the plane of the lateral semicicular canal (SCC) with the head flexed 20 degrees-25 degrees were recorded as a baseline. The lateral SCC was then selectively obliterated near its ampulla. This induced major changes in the VsEPs recorded in response to stimulation of the remaining inner ear in this plane: the first 2 VsEP waves were absent, and only longer latency, smaller amplitude waves were present in response to both clockwise and counterclockwise stimulation. On the other hand, obliteration of the anterior and posterior SCCs and, in addition, destruction of both maculae were without major effects on the first 2 VsEP waves in response to excitatory stimulation. The results confirm that when the head is flexed 20 degrees-25 degrees and stimulated with angular acceleration impulses in the horizontal plane, the major site of initiation of the VsEPs in cats and probably in man is the crista ampullaris of the lateral SCC.

The pattern of inputs from six semicircular canals to neck motoneurons was investigated by stimulating six ampullary nerves electrically and recording intracellular potentials from motoneurons of the rectus capitis dorsalis (RD), the complexus (COMP) and the obliquus capitis caudalis (OCA) muscles at the upper cervical cord of the cat. RD and COMP motoneurons received disynaptic excitation from bilateral anterior and contralateral horizontal ampullary nerves and disynaptic inhibition from bilateral posterior and ipsilateral horizontal ampullary nerves. OCA motoneurons received excitation from ipsilateral vertical and contralateral horizontal ampullary nerves and inhibition from contralateral vertical and ipsilateral horizontal ampullary nerves. Ipsilateral disynaptic inhibitory postsynaptic potentials and contralateral disynaptic excitatory postsynaptic potentials to these motoneurons were mediated by the medial longitudinal fasciculus (MLF) and the other postsynaptic potentials by the extra-MLF pathways. The results indicated that motoneurons of a neck muscle have its own characteristic pattern of inputs from six semicircular canals.

Motor control of different segments of the body with multiple degrees of freedom appears to be coordinated by utilizing preferred axes of motor activity. This hypothesis may also be applied to vestibular control of posture. To explore this question we studied the anatomical relationship between the head and the cervical vertebral column by taking radiographs of the head-neck region in unrestrained alert guinea pigs. We determined that biomechanical constraints contribute to the stereotypical skeletal geometry observed in the resting animal and to a functional segmentation of the head-neck movement apparatus. Subsequent lesion studies of vestibular end organs with quantification of the resulting postural syndromes suggest that the functional segmentation of the cervical vertebral column corresponds to a functional partitioning of vestibular afferents. Our findings also indicate that the sensorimotor transformation mechanisms necessary to convert a given head velocity signal into the appropriate neck motor frame are already embedded in the networks provided by second-order vestibular neurons. Good trim of postural control will be the end result of an appropriate internal representation of the objective vertical.

Floccular influence on excitatory vestibular reflex arcs of anterior semicircular canal origin was examined in the anaesthetized cat. Stimulation of the anterior semicircular canal nerve (ACN) evoked disynaptic excitatory postsynaptic potentials (EPSPs) in all sampled inferior oblique (IO), superior rectus (SR), and biventor cervicis (BIV) muscle motoneurones of the contralateral side. Conditioning stimulus to the flocculus depressed the amplitude of the EPSPs in both IO and SR motoneurones by 50% on the average but not in any BIV motoneurones. The excitatory vestibulo-ocular neurones identified by orthodromic and antidromic responses to stimulation of the ACN and the contralateral IO motoneurone pool, respectively, were classified as VOC (vestibulo-ocular neurones with axons descending to the cervical segment) or VO (vestibulo-ocular proper) neurones on the basis of whether or not they responded antidromically to stimulation of the spinal cord in the C1 segment. All of the VO neurones in the superior vestibular nucleus (n = 19) were inhibited from the flocculus while the activities of three-fourths of the VO neurones (36/48) in the other vestibular nuclei were not suppressed by floccular stimulation. In contrast, none of VOC neurones (n = 49) received floccular inhibition. Besides inhibition, floccular stimulation induced the antidromic or orthodromic responses in some VO and VOC neurones.

In normal subjects transient horizontal body rotation in the dark (vestibular stimulation) elicited a DC potential change with the maximum at Cz. The response appeared to covary with stimulus velocity, which is the most relevant stimulus parameter transferred by the horizontal semicircular canal system. A similar response was obtained during optokinetic stimulation. However, the following findings suggested that the responses are related to 'high' perceptual functions rather than representing a visuo-vestibular evoked cortical potential: (a) the visual response was found independent of whether the subjects perceived, as a task, self-motion in a stationary environment or 'object' motion about the stationary body; its amplitude depended on the subjects' subjective compliance to the task; (b) when presenting various combinations of optokinetic and vestibular stimuli, the response amplitude depended on the subjective and objective intersensory conflict in the combinations; (c) sinusoidal stimulation yielded a negative shift of the DC potential, but the potential was not modulated along with the waxing and waning of stimulus velocity or of the self-motion sensation evoked; (d) patients with loss of vestibular functions showed a similar Cz response during body rotation in the dark.

1. A new concept of endolymph flow in the vertebrate vestibular system is presented. This approach describes quantitatively the flow in the entire system of three semicircular ducts interconnected by the utriculus and the crus commune. This approach is quite distinct from the classical theory in which the labyrinth is generally conceived to consist of three separate duct circuits. 2. The present approach shows the following set of distinct differences to the classical view: (a) In a labyrinth composed of three ducts perpendicular to each other the flow is non-zero in the other ducts when the labyrinth is rotated in the plane of a particular duct. (b) In a labyrinth with two equal ducts and with the duct planes under approximately 73 degrees the flow in one duct is zero when the rotation takes place in the plane of the other duct. Previous measurements of duct angles reflect this value surprisingly well. An obtuse or sharp angle between duct planes can lead to better performance of a particular labyrinth because the "external impulses" in the different ducts may amplify or compensate each other. (c) The behaviour of the flow in the entire labyrinth is a non-linear function of direction or rotation (cf. points (d), (e]. (d) Six time constants for the entire labyrinth can be distinguished (three long, three short); the flow in a particular duct is composed of six terms with these time constants. The composition of this flow and thus the relative importance of the terms depends on the positioning of the labyrinth with respect to the rotation vector. (e) The time constants also depend, for different labyrinths, on a shared influence of the dimensions of the ducts and the elastic properties of all three cupulae. (f) The forces in a particular duct depend also on the amount of motion the fluid will acquire in the other ducts. (g) The sensitivity of a particular duct depends also on the dimensions of the other parts in the vestibular system. 3. Equations for a system consisting of two ducts and for the classical single duct system are also given. Both systems are special cases of the three-duct system. The single duct equations are equivalent with equations given by Oman (1980) and Oman et al. (1987) which include the contribution of a wide utriculus. 4. The present theory of endolymph flow is mainly supported by the outcome of previously performed experiments concerning time constants and rotation of human subjects in different planes.(ABSTRACT TRUNCATED AT 400 WORDS)

1. A mathematical treatment of the flow inside the vertebrate labyrinth is given. The main difference to former theories (e.g. the "torsion pendulum" theory) is that the entire system formed by the three semicircular ducts, interconnected by the crus commune and the utriculus, is considered, instead of a single duct circuit. 2. The theory consists of a geometrical description of a labyrinth rotating in space, the solution of the continuity equation, determination of the initial velocities in all the ducts in a "cupulometry" experiment and derivation of the equation of motion (e.o.m.). 3. Equations for a system consisting of two ducts and for the classical single-duct system are special cases of the three-duct system. 4. Three different methods for the solution of the e.o.m. are described: an analytical one, a Runge-Kutta simulation and an "asymptotic" method. The last method includes approximations of the solution of the e.o.m. on a long and a short time scale. Its advantage is that it gives an insight based on rather manageable formulae. 5. The physiological basis of the presented theory, biological applications and verification are given in a separate paper (Muller & Verhagen, 1988).