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The Vestibular System

Jay M. Goldberg, César Fernández

10.1002/cphy.cp010321

Source: Supplement 3: Handbook of Physiology, The Nervous System, Sensory Processes

Originally published: 1984

Published online: January 2011

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Peripheral Receptor Mechanisms
1.1 Morphology of Vestibular Apparatus
1.2 Mechanisms of Sensory Transduction
1.3 Resting Discharge
1.4 Fiber Caliber and Afferent Response
1.5 Physiology of Semicircular Canals
1.6 Physiology of Otolith Organs
2 Centrifugal Systems
2.1 Efferent System
2.2 Sympathetic Innervation
2.3 Receptor‐Receptor Neurons
3 Vestibular Nuclei
3.1 Functional Organization
3.2 Physiology of Canal‐Related Secondary Neurons
3.3 Physiology of Otolith‐Related Secondary Neurons
3.4 Sensitivity to Nonlabyrinthine Inputs
3.5 Vestibuloocular Reflexes
3.6 Canal‐Related Reflexes
3.7 Otolith‐Related Reflexes
3.8 Barbecue Nystagmus
3.9 Vestibulospinal Relations
3.10 Lateral Vestibulospinal Tract
3.11 Medial Vestibulospinal Tract
3.12 Functional Considerations
3.13 Vestibulocerebellar Relations
3.14 Inputs to Vestibulocerebellum
3.15 Cerebellar Influences on Vestibular Pathways
4 Vestibulocortical Projections
4.1 Cortical Representation
4.2 Thalamic Representation and Brain Stem Pathways
5 Plasticity in Vestibular Pathways
5.1 Compensation from Labyrinthectomy
5.2 Habituation
5.3 Adaptive Control of Vestibuloocular Reflex
Figure 1. Figure 1.

Gross anatomy of membranous labyrinth and its innervation in mammals, c.r., Canal reuniens; e.d., endolymphatic duct; H, P, and S, horizontal, posterior, and superior semicircular canals, respectively; Sa, sacculus; SG, Scarpa's ganglion; U, utriculus; VII, facial nerve. Superior vestibular nerve innervates superior and horizontal canals and utriculus; a small branch (Voigt's anastomosis) runs to the sacculus. Inferior vestibular nerve supplies sacculus and posterior semicircular canal; a small branch (Oort's anastomosis) contains auditory efferents destined for the cochlea.

From Hardy 135
Figure 2. Figure 2.

Structure of crista ampullaris in mammals. A: relation of sensory hair cells to their innervation (bottom) and to the cupula (top). B: ultrastructural organization of type I and type II hair cells and their innervation.

From Wersäll and Bagger‐Sjöbäck 349
Figure 3. Figure 3.

Structural organization of A: mammalian saccular macula; B: utricular macula. Illustrated are regional differences in otolithic membranes and otoconia, in structure and type of hair cells, and in patterns of afferent innervation. Note reversal in morphological polarization of sensory cells in region of each striola.

From Lindeman 189
Figure 4. Figure 4.

Organization of mammalian vestibular nerve. EF, efferent nerve fibers; H.C.A., P.C.A., and S.C.A., horizontal, posterior, and superior canal ampullae, respectively; P.C.N., Sacc.N., and Utric.N., posterior canal, saccular, and utricular nerves, respectively; Sacc., sacculus; S.G., Scarpa's ganglion; Utr., utriculus.

From Gacek 101
Figure 5. Figure 5.

Morphological polarization of sensory cells and polarization maps of mammalian sensory epithelia. A: section of sensory epithelium. Kinocilium (filled bars) and stereocilia (open bars) are seen. Arrow indicates direction of excitatory polarization. B: section through sensory hair bundles showing relative locations of kinocilium and stereocilia. C: sensory cells of each crista ampullaris are all polarized in same direction (arrows). D, E: in each otolith organ the polarization reverses in region of striola (dashed line).

From Lindeman 189
Figure 6. Figure 6.

Resting discharge of two semicircular canal units recorded in squirrel monkey. Though both afferents have a similar firing rate (close to 100 spikes/s), they differ in the regularity of their discharge patterns.

From Goldberg and Fernández 116
Figure 7. Figure 7.

Calculated displacement of cupula and endolymph (in arbitrary units) to A: long‐duration acceleration step; to B: brief, bidirectional head rotation. Below each figure are shown velocity profile (solid line) and acceleration profile (dashed line).
Figure 8. Figure 8.

Calculated response to sinusoidal inputs of the cupula and endolymph based on the torsion‐pendulum model. Log gains and phases with respect to angular head velocity. LF, MF, HF: low‐, medium‐, high‐frequency ranges, respectively. Gains are expressed relative to their value in the MF range; phase leads are negative.

From Melvill‐Jones and Milsum 236
Figure 9. Figure 9.

Response of semicircular canal neurons to velocity trapezoids. Bars, periods of constant angular acceleration or deceleration. A‐B: responses of a regular unit to (A) 5‐s, 60°/s2, and (B) to 40‐s, 7.5°/s2 acceleration steps. C, D: responses of an irregular unit to same stimulus conditions. Note adaptive phenomena.

From Goldberg and Fernández 116
Figure 10. Figure 10.

Responses of semicircular canal neurons to sinusoidal rotations. Data are for a regular (○) and an irregular (•) unit. Log gains and phases (——) predicted from torsion‐pendulum model; curves based on empirical transfer functions, (—–) and (——‐). Gains and phases are expressed relative to angular head velocities and gains are normalized at 0.25 Hz.

From Fernández and Goldberg 81
Figure 11. Figure 11.

Responses of an otolith neuron to linear forces. A: discharge rate vs. angle of static head tilt. Direction of gravity relative to head was changed by sequentially tilting the animal in 45° steps around either pitch, •, or roll, ○, axes. At each position a 10‐s sample of activity was taken. Positive tilt angles correspond to forward pitches and ipsilateral rolls. Solid and dashed lines are best fitting trigonometric functions for pitch and roll points, respectively. From the data, a functional polarization vector was calculated. B: response to centrifugal force trapezoids with a peak force of 1.23 g; bars, periods of force transition. Force was directed parallel, •, and antiparallel, ○, to unit's polarization vector. Note asymmetry in corresponding excitatory and inhibitory responses. C: force‐response relations, based on data in A and B. The force, xlg, is calculated from Equation 8 and the solid curve is based on Equation 7. There is good agreement between the force‐response relations for static tilts, ○, and for centrifugal force, •.

From Fernández and Goldberg 82
Figure 12. Figure 12.

Origin and course of efferent vestibular system in cat. A: distribution of neurons retrogradely labeled after unilateral horseradish peroxidase injection into labyrinth of a newborn animal. Efferent neurons are seen in horizontal plan view. Top of diagram, rostral direction. Each dot, •, approximately 5 labeled neurons; bar, 1 mm; SVN, LVN, DVN, MVN: superior, lateral, descending, and medial vestibular nuclei, respectively; VI, abducens nucleus. B: frontal section (see arrow in part A) from same animal. Each dot, •, one labeled neuron; Sp. Tr. V, trigeminal spinal tract; V, spinal trigeminal nucleus; VII, facial nerve; MLF, medial longitudinal fasciculus; BC, brachium conjunctivum; all other parts identified under A. C: intramedullary course of efferent pathways based on acetylcholinesterase histochemistry. RB, restiform body; CN, cochlear nuclei; all other parts identified under A and B.
Figure 13. Figure 13.

Afferent and efferent innervation in mammalian sensory epithelium. One efferent nerve fiber is shown making both pre‐ and postsynaptic contacts by means of vesiculated boutons, VB. BM, basement membrane; C, afferent nerve chalice; SB, synaptic bar.

From Smith and Rasmussen 311
Figure 14. Figure 14.

Topography and cytoarchitecture of vestibular nuclear complex in the cat as seen in transverse sections. B.c., brachium conjunctivum; C.r., restiform body; D., descending vestibular nucleus; L., lateral vestibular nucleus; M., medial vestibular nucleus; N.cu.e., external cuneate nucleus; N.i.n. VIII, interstitial nucleus of vestibular nerve; N.V, VII, VIII, IX, XII, cranial nerves; p.h., nucleus praepositus hypoglossi; S., superior vestibular nucleus; Tr.sp. V, spinal tract of trigeminal nerve; V, VI, VII, X, XII, motor cranial nerve nuclei; f, g, l, Sv, x, y, z, small cell groups.

From Brodal and Pompeiano 45, by permission of Cambridge Univ. Press
Figure 15. Figure 15.

Responses of several secondary, horizontal‐canal‐related neurons (denoted by different symbols) to sinusoidal rotations. Log gains and phase leads are plotted with respect to angular head velocity. Dashed lines, expected behavior of the torsion pendulum model with time constants, T1.

From Melvill‐Jones and Milsum 236
Figure 16. Figure 16.

A: field potentials recorded from vestibular nuclei upon electrical stimulation of ipsilateral vestibular nerve. Upward is negative. Calibrations: 1 ms and 1 mV. B: organization of commissural pathways. Inhibitory neurons are shown as filled circles. Dashed line indicates midline. V.n., vestibular nerve; Ik and It, kinetic and tonic type I neurons, respectively.

A: from Precht and Shimazu 269; B: from Precht 380
Figure 17. Figure 17.

Eye‐head coordination in intact and labyrinthectomized monkey. A: normal monkey. The animal fixates a new visual target by making an eye movement, E, and a slower head movement, H. During head movement the eyes move backward relative to the head so that the gaze, G, remains constant. B, C: 40 and 120 days, respectively, after labyrinthectomy.

From Dichgans et al. 66
Figure 18. Figure 18.

Eye movements evoked in cat by electrical stimulation of ampullary nerves. A‐C: unilateral stimulation. D‐F: bilateral stimulation. Eyes are seen from the front; arrows, direction of movement. Primary excitation of individual extraocular muscles in black; secondary excitation indicated by stippling.

From Cohen et al. 57
Figure 19. Figure 19.

Block diagram of vestibuloocular reflex. It is assumed that the head moves through an angle θ = sin ωt. Output of each component is shown below the corresponding arrow; gains and phase leads with respect to angular head position are in parentheses.
Figure 20. Figure 20.

Dynamics of various components of vestibuloocular reflex expressed as phase leads relative to angular head position (θh). Upper border of figure represents the physical stimulus (head acceleration, Θh), which leads θh by 180°. Top curve, Rv shows phase lag, α, contributed by the semicircular canals. Middle curve, EOM, includes the phase lag, γ, introduced by extraocular muscles and orbital mechanics. Bottom curve, θe/h, represents actual performance of the reflex and requires that a phase lag, β, be inserted by the central pathways.

From Skavenski and Robinson 306
Figure 21. Figure 21.

Rhythmic changes in intracellular potential of abducens motoneuron and motor nerve discharges during nystagmus. Responses during high‐frequency electrical stimulation to A: left vestibular nerve; B: right vestibular nerve. Top traces in A and B, records from a motoneuron, on right side; middle traces, from left abducens nerve; bottom traces, from right abducens nerve. Upward deflection indicates positivity.

From Maeda et al. 215
Figure 22. Figure 22.

Uncoordinated eye movements elicited in anesthetized cat by focal stimulation of corresponding part of A: left utricular macula, or B: left saccular macula. Eyes are viewed from the front. Arrows, motions of the right, Rt, and left, Lt, eyes. A.L., A.M., M.L., P.L., and P.M. refer to anterolateral, anteromedial, mediolateral, posterolateral, and posteromedial parts of utricular macula, respectively; SUP. and INF., superior and inferior parts of saccular macula, respectively.

A: adapted from Fluur and Mellström 90; B: adapted from Fluur and Mellström 91
Figure 23. Figure 23.

Connections between ipsilateral and contralateral ampullae and neck extensor motoneurons in the cat. A, H, P are anterior, horizontal, and posterior ampullae; VN, vestibular nuclei. Inhibitory neurons and their terminals shown in filled symbols, excitatory neurons in open symbols.

From Wilson and Maeda 357
Figure 24. Figure 24.

Vestibulocerebellar relations in cat. A: vestibular nerve is shown to send projections to flocculus, Flocc., to dorsal, Pfl.d., and ventral, Pfl.v., paraflocculus; to uvula, IX(Uv.), and nodulus, X(Nod.), to lingula, I; and to parvocellular region, p, of lateral cerebellar nucleus, Nl. B: secondary vestibular projection from medial, M, descending, D, superior, S, and lateral, L, vestibular nuclei, and x group to flocculus, Flocc.; Nod., nodulus; Uv., uvula; and N. fast., fastigial nucleus. C: projections from vestibulocerebellum to vestibular nuclei; f and x, small cell groups; other nuclei identified in part B.

A: from Brodal and Høivik 44; B: from Brodal 42; C: from Angaut and Brodal 17


Figure 1.

Gross anatomy of membranous labyrinth and its innervation in mammals, c.r., Canal reuniens; e.d., endolymphatic duct; H, P, and S, horizontal, posterior, and superior semicircular canals, respectively; Sa, sacculus; SG, Scarpa's ganglion; U, utriculus; VII, facial nerve. Superior vestibular nerve innervates superior and horizontal canals and utriculus; a small branch (Voigt's anastomosis) runs to the sacculus. Inferior vestibular nerve supplies sacculus and posterior semicircular canal; a small branch (Oort's anastomosis) contains auditory efferents destined for the cochlea.

From Hardy 135


Figure 2.

Structure of crista ampullaris in mammals. A: relation of sensory hair cells to their innervation (bottom) and to the cupula (top). B: ultrastructural organization of type I and type II hair cells and their innervation.

From Wersäll and Bagger‐Sjöbäck 349


Figure 3.

Structural organization of A: mammalian saccular macula; B: utricular macula. Illustrated are regional differences in otolithic membranes and otoconia, in structure and type of hair cells, and in patterns of afferent innervation. Note reversal in morphological polarization of sensory cells in region of each striola.

From Lindeman 189


Figure 4.

Organization of mammalian vestibular nerve. EF, efferent nerve fibers; H.C.A., P.C.A., and S.C.A., horizontal, posterior, and superior canal ampullae, respectively; P.C.N., Sacc.N., and Utric.N., posterior canal, saccular, and utricular nerves, respectively; Sacc., sacculus; S.G., Scarpa's ganglion; Utr., utriculus.

From Gacek 101


Figure 5.

Morphological polarization of sensory cells and polarization maps of mammalian sensory epithelia. A: section of sensory epithelium. Kinocilium (filled bars) and stereocilia (open bars) are seen. Arrow indicates direction of excitatory polarization. B: section through sensory hair bundles showing relative locations of kinocilium and stereocilia. C: sensory cells of each crista ampullaris are all polarized in same direction (arrows). D, E: in each otolith organ the polarization reverses in region of striola (dashed line).

From Lindeman 189


Figure 6.

Resting discharge of two semicircular canal units recorded in squirrel monkey. Though both afferents have a similar firing rate (close to 100 spikes/s), they differ in the regularity of their discharge patterns.

From Goldberg and Fernández 116


Figure 7.

Calculated displacement of cupula and endolymph (in arbitrary units) to A: long‐duration acceleration step; to B: brief, bidirectional head rotation. Below each figure are shown velocity profile (solid line) and acceleration profile (dashed line).



Figure 8.

Calculated response to sinusoidal inputs of the cupula and endolymph based on the torsion‐pendulum model. Log gains and phases with respect to angular head velocity. LF, MF, HF: low‐, medium‐, high‐frequency ranges, respectively. Gains are expressed relative to their value in the MF range; phase leads are negative.

From Melvill‐Jones and Milsum 236


Figure 9.

Response of semicircular canal neurons to velocity trapezoids. Bars, periods of constant angular acceleration or deceleration. A‐B: responses of a regular unit to (A) 5‐s, 60°/s2, and (B) to 40‐s, 7.5°/s2 acceleration steps. C, D: responses of an irregular unit to same stimulus conditions. Note adaptive phenomena.

From Goldberg and Fernández 116


Figure 10.

Responses of semicircular canal neurons to sinusoidal rotations. Data are for a regular (○) and an irregular (•) unit. Log gains and phases (——) predicted from torsion‐pendulum model; curves based on empirical transfer functions, (—–) and (——‐). Gains and phases are expressed relative to angular head velocities and gains are normalized at 0.25 Hz.

From Fernández and Goldberg 81


Figure 11.

Responses of an otolith neuron to linear forces. A: discharge rate vs. angle of static head tilt. Direction of gravity relative to head was changed by sequentially tilting the animal in 45° steps around either pitch, •, or roll, ○, axes. At each position a 10‐s sample of activity was taken. Positive tilt angles correspond to forward pitches and ipsilateral rolls. Solid and dashed lines are best fitting trigonometric functions for pitch and roll points, respectively. From the data, a functional polarization vector was calculated. B: response to centrifugal force trapezoids with a peak force of 1.23 g; bars, periods of force transition. Force was directed parallel, •, and antiparallel, ○, to unit's polarization vector. Note asymmetry in corresponding excitatory and inhibitory responses. C: force‐response relations, based on data in A and B. The force, xlg, is calculated from Equation 8 and the solid curve is based on Equation 7. There is good agreement between the force‐response relations for static tilts, ○, and for centrifugal force, •.

From Fernández and Goldberg 82


Figure 12.

Origin and course of efferent vestibular system in cat. A: distribution of neurons retrogradely labeled after unilateral horseradish peroxidase injection into labyrinth of a newborn animal. Efferent neurons are seen in horizontal plan view. Top of diagram, rostral direction. Each dot, •, approximately 5 labeled neurons; bar, 1 mm; SVN, LVN, DVN, MVN: superior, lateral, descending, and medial vestibular nuclei, respectively; VI, abducens nucleus. B: frontal section (see arrow in part A) from same animal. Each dot, •, one labeled neuron; Sp. Tr. V, trigeminal spinal tract; V, spinal trigeminal nucleus; VII, facial nerve; MLF, medial longitudinal fasciculus; BC, brachium conjunctivum; all other parts identified under A. C: intramedullary course of efferent pathways based on acetylcholinesterase histochemistry. RB, restiform body; CN, cochlear nuclei; all other parts identified under A and B.



Figure 13.

Afferent and efferent innervation in mammalian sensory epithelium. One efferent nerve fiber is shown making both pre‐ and postsynaptic contacts by means of vesiculated boutons, VB. BM, basement membrane; C, afferent nerve chalice; SB, synaptic bar.

From Smith and Rasmussen 311


Figure 14.

Topography and cytoarchitecture of vestibular nuclear complex in the cat as seen in transverse sections. B.c., brachium conjunctivum; C.r., restiform body; D., descending vestibular nucleus; L., lateral vestibular nucleus; M., medial vestibular nucleus; N.cu.e., external cuneate nucleus; N.i.n. VIII, interstitial nucleus of vestibular nerve; N.V, VII, VIII, IX, XII, cranial nerves; p.h., nucleus praepositus hypoglossi; S., superior vestibular nucleus; Tr.sp. V, spinal tract of trigeminal nerve; V, VI, VII, X, XII, motor cranial nerve nuclei; f, g, l, Sv, x, y, z, small cell groups.

From Brodal and Pompeiano 45, by permission of Cambridge Univ. Press


Figure 15.

Responses of several secondary, horizontal‐canal‐related neurons (denoted by different symbols) to sinusoidal rotations. Log gains and phase leads are plotted with respect to angular head velocity. Dashed lines, expected behavior of the torsion pendulum model with time constants, T1.

From Melvill‐Jones and Milsum 236


Figure 16.

A: field potentials recorded from vestibular nuclei upon electrical stimulation of ipsilateral vestibular nerve. Upward is negative. Calibrations: 1 ms and 1 mV. B: organization of commissural pathways. Inhibitory neurons are shown as filled circles. Dashed line indicates midline. V.n., vestibular nerve; Ik and It, kinetic and tonic type I neurons, respectively.

A: from Precht and Shimazu 269; B: from Precht 380


Figure 17.

Eye‐head coordination in intact and labyrinthectomized monkey. A: normal monkey. The animal fixates a new visual target by making an eye movement, E, and a slower head movement, H. During head movement the eyes move backward relative to the head so that the gaze, G, remains constant. B, C: 40 and 120 days, respectively, after labyrinthectomy.

From Dichgans et al. 66


Figure 18.

Eye movements evoked in cat by electrical stimulation of ampullary nerves. A‐C: unilateral stimulation. D‐F: bilateral stimulation. Eyes are seen from the front; arrows, direction of movement. Primary excitation of individual extraocular muscles in black; secondary excitation indicated by stippling.

From Cohen et al. 57


Figure 19.

Block diagram of vestibuloocular reflex. It is assumed that the head moves through an angle θ = sin ωt. Output of each component is shown below the corresponding arrow; gains and phase leads with respect to angular head position are in parentheses.



Figure 20.

Dynamics of various components of vestibuloocular reflex expressed as phase leads relative to angular head position (θh). Upper border of figure represents the physical stimulus (head acceleration, Θh), which leads θh by 180°. Top curve, Rv shows phase lag, α, contributed by the semicircular canals. Middle curve, EOM, includes the phase lag, γ, introduced by extraocular muscles and orbital mechanics. Bottom curve, θe/h, represents actual performance of the reflex and requires that a phase lag, β, be inserted by the central pathways.

From Skavenski and Robinson 306


Figure 21.

Rhythmic changes in intracellular potential of abducens motoneuron and motor nerve discharges during nystagmus. Responses during high‐frequency electrical stimulation to A: left vestibular nerve; B: right vestibular nerve. Top traces in A and B, records from a motoneuron, on right side; middle traces, from left abducens nerve; bottom traces, from right abducens nerve. Upward deflection indicates positivity.

From Maeda et al. 215


Figure 22.

Uncoordinated eye movements elicited in anesthetized cat by focal stimulation of corresponding part of A: left utricular macula, or B: left saccular macula. Eyes are viewed from the front. Arrows, motions of the right, Rt, and left, Lt, eyes. A.L., A.M., M.L., P.L., and P.M. refer to anterolateral, anteromedial, mediolateral, posterolateral, and posteromedial parts of utricular macula, respectively; SUP. and INF., superior and inferior parts of saccular macula, respectively.

A: adapted from Fluur and Mellström 90; B: adapted from Fluur and Mellström 91


Figure 23.

Connections between ipsilateral and contralateral ampullae and neck extensor motoneurons in the cat. A, H, P are anterior, horizontal, and posterior ampullae; VN, vestibular nuclei. Inhibitory neurons and their terminals shown in filled symbols, excitatory neurons in open symbols.

From Wilson and Maeda 357


Figure 24.

Vestibulocerebellar relations in cat. A: vestibular nerve is shown to send projections to flocculus, Flocc., to dorsal, Pfl.d., and ventral, Pfl.v., paraflocculus; to uvula, IX(Uv.), and nodulus, X(Nod.), to lingula, I; and to parvocellular region, p, of lateral cerebellar nucleus, Nl. B: secondary vestibular projection from medial, M, descending, D, superior, S, and lateral, L, vestibular nuclei, and x group to flocculus, Flocc.; Nod., nodulus; Uv., uvula; and N. fast., fastigial nucleus. C: projections from vestibulocerebellum to vestibular nuclei; f and x, small cell groups; other nuclei identified in part B.

A: from Brodal and Høivik 44; B: from Brodal 42; C: from Angaut and Brodal 17
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Article Title: The Vestibular System
 

How to Cite

Jay M. Goldberg, César Fernández. The Vestibular System. Compr Physiol 2011, Supplement 3: Handbook of Physiology, The Nervous System, Sensory Processes: 977-1022. First published in print 1984. doi: 10.1002/cphy.cp010321