Anat 6.8 Auditory Pathways_Bayaoa
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Anatomy 6.8
February 21, 2012 Dra. Bayaoa
Auditory Pathways
Outline I. Auditory and Vestibular system II. Auditory System A. External Ear B. Middle Ear C. Inner Ear III. Central Auditory Pathway IV. Vestibular System A. Inner Ear (Labyrinth) B. Vestibular Functions of the ear Macula Objectives: • Define function of the cochlear system and receptors for hearing • Tracing the pathway of sound from the environment to the sense organ for hearing • Describing the origin, course, and termination of the cochlear nerve • Tracing the central auditory pathway from the organ of Corti until the nerve impulse reaches the auditory cortex • Role of the olivocochlear bundle of Rasmussen in sound perception • Pathways of auditory reflexes • Differentiating between conductive and sensory deafness • Explaining tinnitus, Rinne’s & Weber’s test • Function of the vestibular system • Parts of Static labyrinth vs kinetic labyrinth • Location of the receptors for vestibular control • Origin, course, and termination of the vestibular nerve • Describing the role of: MLF, medial vestibulospinal tract, lateral vestibulospinal tract in the maintenance of equilibrium • Connections of the vestibular apparatus with the cerebellum • Define: vertigo and nystagmus • Tests for vestibular function • Clinical manifestations of disturbances in vestibular apparatus
I. AUDITORY AND VESTIBULAR SYSTEM CN VIII/ Vestibulocochlear nerve • Has two distinct divisions 1. Vestibular – position and movement of head; for equilibrium 2. Cochlear – mediates auditory function • Grossly; it is divided into external ear, middle ear and inner ear • Stimulus: SOUND o Sinusoidal waves of air molecules o Frequency of waves (in hertz) -‐> PITCH of sound o Amplitude of each wave (in decibels) – LOUNDESS of sound o The human ear can detect sound frequencies from 20 to 20, 000 Hz and 1-‐120 db II. AUDITORY SYSTEM A. External Ear Consists of • Auricle/pinna/ear lobe • External auditory meatus ( ear canal) • Tympanic Membrane (ear drum; separates the external ear from the middle ear) Sound waves enter the external auditory meatus -‐> impinge on the tympanic membrane -‐> tympanic membrane vibrates and transmits sound into the middle ear B. Middle Ear • Lies in the petrous part of temporal bone • Transmits the vibrations of the tympanic membrane to the inner ear RG, SOPIE, KEIFER, TASIE, FILLE, ELIZ, MAY, JANNA J
• Air-‐filled cavity • Two parts: o Tympanic cavity proper o Attic/ epitympanic recess
Figure 1. Structures of the middle ear (Coronal Oblique Section)
• Consists of the three auditory ossicles (MIS) 1. Malleus (hammer) – attaches to the inner aspect of the tympanic membrane 2. Incus (anvil) – receives vibration from the malleus via diarthrodial joint; articulates with stapes 3. Stapes (stirrup) – sits on the membrane of the foramen ovale (oval window) through which vibration is transmitted to the fluid perilymph of the inner ear. • The 3 auditory ossicles serve as an amplifier and as an impedance-‐ matching device that decreases the amount of energy lost by by the sound waves in going from the air to the fluid in the inner ear • Eustachian tube o From the cavity of the middle ear to the posterior nasopharynx o Equalizes air pressure inside and outside tympanic membrane • Contains two muscles: 1. Stapedius -‐ innervated by CN VII (facial) • pulls on stapes and causes tightening of the membrane of the oval window 2. Tensor tympani -‐ innervated by CN V (mandibular branch of trigeminal nerve) § pulls on malleus and causes the tightening of the tympanic membrane • The two muscles contract to decrease transmission of vibrational energy from external to inner ear to protect the inner ear from loud sounds (attenuation reflex)
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• Hyperacousis -‐ sounds perceived as uncomfortable and louder. May be experienced by person with paralysis of these 2 muscles or of the stapedius alone (as in Bell’s facial palsy) C. Inner Ear • Two components: o Cochlea -‐ organ for hearing, where cochlear nerve attaches o Vestibular apparatus (3 semicircular canals, utricle and saccule) – where vestibular nerve attaches • The oval window/foramen ovale, to which the stapes communicates, opens to the vestibule portion which contains perilymph • Bony Labyrinth (contains perilymph) is made up of the: o Cochlea o Semicircular canals o Vestibule • Membranous labyrinth (with endolymph) is found within the bony labyrinth and is made up of the: o cochlear duct (in bony cochlea) o utricle and saccule (in vestibule) o semicircular ducts (in semicircular canals) 1. Cochlea
Figure 2. The cochlea
• Resembles a snail • Contains helicotrema which is the apical connection between the scala vestibuli and scala tympani. It is also the base of cochlea • Turns at its conical central axis called modiolus. • 2 and ¾ turns • Consists of three parallel filled channels: o Scala vestibuli (w/ perilymph) o Scala tympani (w/ perilymph) o Scala media or cochlear duct (w/ endolymph) *Perilymph is similar to CSF while endolymph is similar to intracellular fluid. • Vestibular membrane divides scala vestibuli into another space called scala media or cochlear duct RG, SOPIE, KEIFER, TASIE, FILLE, ELIZ, MAY, JANNA J
• Contains a spiral lamina (ridge of bone dividing scala vestibuli and scala tympani) • Scala media/cochlear duct -‐ a membranous labyrinth within the cochlea, completes the separation of scala vestibuli and scala tympani • Basilar membrane -‐ where the cochlea sits. It is narrower at base than at the apex. 2. Organ of Corti • Within the cochlear duct/scala media • Generates action potentials in the bipolar neurons of the spiral ganglion of the cochlear division of the vestibulocochlear nerve when hair cells vibrate due to sound. • Functions as an audiofrequency analyzer/organizer o Highest tones stimulate hair cells near the basal cochlea which contains the narrowest segment of basilar membrane o Lowest tones stimulate hair cells in the apical portion of the cochlea which contains the widest segment of the basilar membrane o Intermediate tones stimulate hair cells on the intermediate portion of the basilar membrane • Consists of two receptor cells: a. Inner hair cells • Auditory receptor cells • Base o attached to basilar membrane o synapses with dendrites of spiral ganglion cells (10 spiral ganglion cells innervate 1 hair cell and its axons form the cochlear division of CN VIII) o inner hair cells + spiral ganglion = produce frequency-‐ dependent responses to sound • Apex o contains stereocilia which lie just below the tectorial membrane o Stereocilia touch the tectorial membrane when sound waves enter the cochlea à bending of the stereocilia à open ionic channels à causes changes of potential in hair cell membrane
Figure 3. Organ of Corti
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(N1) spiral ganglion
b.Outer Hair cell • Stereocilia is found embedded in the tectorial membrane • Possesses contractile properties • Controls sensory response properties of the organ of Corti by regulating apposition of tectorial membrane to inner hair cells III. CENTRAL AUDITORY PATHWAY 1. Receptor (Organ of Corti) 2. N1 (spiral ganglion) 3. N2 (ventral and dorsal cochlear nuclei) *fibers form 3 acoustic striae (in bold letters) a. Dorsal cochlear nucleus à dorsal acoustic stria à (most of them cross to join) contralateral lateral lemniscus b.Ventral cochlear nucleus à intermediate acoustic stria (course similar to that of the dorsal acoustic stria) à contralateral lateral lemniscus • Dorsal and intermediate acoustic striae constitute the central monaural auditory pathway, carrying information about the frequency of auditory signals. c. Ventral cochlear nucleus à ventral acoustic stria à terminate in ipsilateral and contralateral nuclei of the trapezoid body and superior olivary nuclei à ipsilateral and contralateral lemnisci • Ventral acoustric stria forms a binaural pathway which analyzes the location of origin, or direction, of auditory stimuli. 4. N3 (inferior colliculus) • Sends axons to medial geniculate nucleus through the brachium of the inferior colliculus 5. N4 (medial geniculate body) • Final sensory relay station of hearing pathway • Special sensory nuclei of the thalamus • Auditory radiation through the sublenticular portion of the internal capsule will go to the transverse temporal gyri (of Heschl) and to planum temporal 6. Primary auditory cortex • Heschl’s gyrus or anterior transverse temporal gyrus or • Brodmann areas 41 and 42 *When impulses reach areas 41 and 42, sound is heard. It is at area 22 (auditory association area) where interpretation occurs. Commissural fibers exist between the following nuclei in the auditory pathway: • Superior olivary nuclei • Nuclei of trapezoid body • Nuclei of lateral lemniscus • Inferior colliculi
RG, SOPIE, KEIFER, TASIE, FILLE, ELIZ, MAY, JANNA J
(N2)
dorsal cochlear nucleus (ipsilateral)
ventral cochlear nucleus (ipsilateral)
Dorsal acoustic stria Intermediate acoustic stria Ventral acoustic stria
trapezoid nuclei superior olivary nuclei (contralateral and ipsilateral) Lateral Lemniscus Lateral Lemniscus (contralateral) (ipsilateral) (N3) Inferior colliculus Inferior colliculus (N4) MGB MGB Primary auditory complex (BA 41 & 42)
Tonotopic Representation in the Auditory System Low Tones High Tones Organ of Corti Upper Lower Cochlear Nuclei Ventral Dorsal MGB Lateral Medial Heschl’s gyrus Anterolateral Posteromedial A. Efferent Transmission • Heschl’s gyrus à medial geniculate body à inferior colliculus à superior olive à cochlear nuclei à organ of Corti • Efferent cochlear bundle/olivocochlear bundle = terminates at organ of Corti where fibers end in synaptic relationship to hair cells • Feedback mechanism to sharpen tone of perception • For modulation, suppression, selection of impulses arising in organ of Corti B. Auditory Reflexes • Involuntary responses to sound mediated by branches of main auditory pathway 1. Audiomotor reflexes o High-‐intensity sound stimulus à stapedius and tensor tympani muscles of the ossicles contract (stapedius pulls the stapes (stirrup) of the middle ear away from the oval window of the cochlea and the tensor tympani muscle pulls the malleus (hammer) away from ear drum) à diminish vibration of middle ear ossicles à decreased transmission of vibrational energy to the cochlea o Impulse goes to cochlear nuclei à R and L superior olive à reticular formation à CN V (for tensor tympani ms) and CN VII (for stapedius ms)
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• For the succeeding pathways, follow: o Inferior colliculus à superior colliculus à tectospinal tract à lower motor neurons in the brain stem and cervical spinal cord, and supply the oculomotor system and muscles of the head and neck that respond to sound 2. Head and neck turning in response to sound 3. General acoustic muscle reflex o Generalized jerking of the body in response to a loud, sudden sound 4. Auditory-‐oculogyric reflex o Deviation of the eyes in the direction of the sound 5. Auditory palpebral reflex o Blinking of the eyelids in response to a loud sound 6. Cochleopupillary reflex o Dilation of the pupils in response to a loud sound C. Hearing Defects 1.Nerve deafness • Destruction of cochlear portion of CN VIII • Lesion of lateral lemniscus = partial deafness only since pathways are bilateral 2.Occlusion deafness • When conduction of sound is impaired before receptor is reached • Due to impacted cerumen, otitis media (infection in middle ear) or otosclerosis (arthritis of auditory ossicles) 3.Tinnitus • hissing, roaring, buzzing, humming sounds due to acoustic neuroma, streptomycin, aspirin • may lead to nerve deafness E. Screening for hearing loss 1. Weber test • Reflects conduction loss in the ipsilateral ear o Conduction problem of the middle ear (incus, malleus, stapes, and eustachian tube) masks the ambient noise of the room, making the sound appear louder o The well-‐functioning inner ear (cochlea with its basilar membrane) picks the sound up via the bones of the skull causing it to be perceived as a louder sound than in the affected/abnormal ear (ipsilateral sensorineural hearing heard as louder) • Patient with unilateral conductive hearing loss would hear the tuning fork loudest in the affected ear • Normal Weber = sound is loudest in midline 2. Rinne Test • Used in cases of unilateral hearing loss and establishes which ear has the greater bone conduction by comparing perception of sounds transmitted by air conduction to those transmitted by bone conduction (mastoid process) . o A "positive" result indicates the healthy state, in contrast to many other medical tests; so avoid using the term 'positive' or 'negative', and simply state if the test was normal or abnormal, to avoid confusion. o Normal / positive Rinne: air conduction > bone conduction • Conductive deafness in Weber and Rinne: o Sound lateralizes/is louder in affected ear o In Weber: abnormality in the middle ear structures leads to increased sound perception in abnormal ear
RG, SOPIE, KEIFER, TASIE, FILLE, ELIZ, MAY, JANNA J
o In Rinne: perceptions by bone conduction (via undamaged inner ear) is enhanced compared to air conduction (because of damaged middle ear) • Sensorineural deafness in Weber and Rinne: o Sound is louder in normal ear (Weber) o Bone conduction is ineffective in stimulating damaged nerve o Both bone conduction and air conduction are diminished but air conduction is slightly > bone conduction (Rinne) IV. VESTIBULAR SYSTEM Functions: • Maintenance of body balance • Coordination of eyes, head and body movement • Permits eyes to remain fixed on a point in space as the head moves Consists of: • Receptors located in the inner ear • vestibular hair cells • Peripheral nerves of the vestibular division of CN VIII • Central connections that analyze information about the position and movement of head in space A. Inner Ear (Labyrinth) 1. Bony Labyrinth • Consists of the cochlea, vestibule, semicircular canals • Series of interconnected cavities in the petrous portion of temporal bone • Contains perilymph which fills the space between the bony and the membranous labyrinth
2. Vestibular/Membranous Labyrinth • Consists of the saccule, utricle, semicircular ducts, cochlear duct • Where the peripheral receptors of the vestibular system, the vestibular hair cells, are located • Contains endolymph • 2 swellings within the vestibule o Utricle o Saccule • 3 semicircular canals within the vestibule o Anterior o Lateral/Horizontal o Posterior
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Vestibular Hair Cells • Peripheral receptors of the vestibular system • Reside inside specialized receptor areas within the membranous labyrinth • Sense both dynamic and static functions Dynamic Function: detects linear (translational) and angular (rotational) motion of head in space o Mediated by the semicircular ducts Static Function: detection of position (tilt) of the head o Mediated by the utricle • The apical portion contains numerous long, rigid, unbranched stereocilia, and on one side, a single kinocilium o arranged in rows of increasing length B. Vestibular Functions of the Ear • Movement of the head causes movement of the endolymph and likewise movement of the otolithic membrane (over the macula) and cupula (over the crista ampullaris). This causes bending of the stereocilia. 1. If Hair bundle/ stereocilia bends toward the kinocilium • Tip links (very small strand of protein that connect the + stereocilia) are pulled à cation channels open à influx of K + ions à depolarization of hair cells à opening of Ca channels near base of the cell à stimulates release of neurotransmitter 2. If hair bundle/ stereocilia bends away from the kinocilium • Causes tip links to be slack à closure of apical cation channels + à hyperpolarization à closure of Ca channels à reduce neurotransmitter release
C. Macula
• Specialized receptor region found in the utricle and saccule • The hair cells within it respond to linear acceleration, gravity and tilt of the head • Contains hair cells that synapse with the peripheral nerves/ distal branches of the vestibular ganglion cells • The tips of the stereocilia and kinocilium are embedded in thick, gelatinous layer of proteoglycans called the otolithic membrane, the outer part of which is filled with calcified structures (calcium carbonate) called otoliths (or otoconia) which increases the specific gravity of the otolithic membrane to about twice that of the endolymph. • otolithic membranes move when subjected to acceleration. • 2 types: macula sacculi and macula utriculi o Macula sacculi (lies in the floor of the saccule) lies in a plane perpendicular to macula utriculi (occupies the lateral wall of the utricle), but both are similar histologically. Utricle: located on the floor of the vestibule Saccule: medial wall • The arrangement of hair cells in the macula creates a response to acceleration in any direction. 1. Striola
Figure 6. Striola Figure 4.Movement of stereocilia RG, SOPIE, KEIFER, TASIE, FILLE, ELIZ, MAY, JANNA J
Figure 5. Macula
• Specialized strip through the middle of the macula o Curved equatorial line
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• Defines the site of change in orientation of the hair cells • Zone that runs the length of macula of both utricle and saccule • Because striola curves through the macula, hair cells are polarized in different directions o Utricular and saccular hair cells are directionally sensitive to a wide variety of head positions and linear movements 2. Macula Utricli • Responds to changes in head position with respect to gravity/ tilt and to earth-‐horizontal linear acceleration • Each hair cell is oriented with the kinocilium toward the striola o hair cells of either sides of striola are organized as mirror images o horizontal acceleration depolarizes one sector of the macula regardless of the direction of movement • hair cells are polarized toward the stiola 3. Macula Sacculi • Similar in structure with that of the macula utriculi • Responds to gravitational pull • Oriented vertically, approximately in a parasagittal plane. • The kinocilia are oriented away from striola • Linear acceleration in the vertical direction stimulates the macula o Occurs in response to gravity or against gravity (e.g. acceleration or deceleration in an elevator) • Hair cells are polarized away from the striola D. Semicircular Canals • Anterior (superior) -‐ Kinocilia faces away from the utricle • Lateral (horizontal) -‐ Kinocilia faces the utricle • Posterior -‐ Kinocilia faces away from the utricle • Each semicircular canal has an enlarged end, called the ampulla • Within the ampulla is the ampullary crest or crista ampullaris, a ridge that bears hair cells like those of the maculae
o React to rotational acceleration or angular movement (kinetic equilibrium) o Covered with a gelatinous capsule, the cupula § Extends almost to the roof of the ampulla § Has the same specific gravity as the endolymph, therefore, it cannot sense the effect of gravity RG, SOPIE, KEIFER, TASIE, FILLE, ELIZ, MAY, JANNA J
When the head undergoes angular acceleration, the viscous endolymph in the semicircular ducts pushes in the cupula o Distortion of the cupula evokes a receptor potential in the hair cells of the ampullary crest, à alters the level of activity in the peripheral fibers of CN VIII, innervating the hair cells o The vestibular nerve fibers to each duct respond with an increase in impulse frequency to rotation in one direction and with a decrease in impulse frequency to rotation in the opposite direction • Lodged within the canals are the semicircular ducts : 1. Superior semicircular canal o Vertical o Perpendicular to the long axis of the petrous bone 2. Posterior semicircular canal o Vertical o Parallel with the long axis of the petrous bone 3. Lateral semicircular canal o Horizontal o Lies in the medial wall of the aditus to the mastoid antrum above the facial nerve canal •
Figure 8. Movement of the cristae ampullaris
E. Vestibular Pathways • Afferent fibers of the vestibular nerve have their cell bodies in the vestibular ganglion (of Scarpa) • Axons of bipolar cells of the vestibular ganglion pass through the internal auditory canal and reach the upper medulla in company with the cochlear nerve o Most of the fibers of the vestibular nerve bifurcate into ascending and descending branches and terminate in the vestibular nuclei, which are clustered in the lateral floor of the fourth ventricle o Inferior vestibular nucleus (descending spinal) § Receives input from the semicircular ducts and from the utricle and the saccule § Nucleus projects into the ascending medial longitudinal fasciculus (MLF) o Superior vestibular nucleus (of Bechterew) § Receives input chiefly from the cristae of semicircular canals § Neurons project into the ascending part of the MLF, where they participate in vestibuloocular reflexes o Medial vestibular nucleus (of Schwalbe) § Receives input chiefly from the cristae of semicircular canals § Neurons project into the ascending part of the MLF, where they participate in vestibuloocular reflexes o Lateral vestibular nucleus (of Deiter) § Receives input chiefly from the macula of the utricle § Neurons project to the ascending portion of the MLF and to the spinal cord through the lateral vestibulospinal tract
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• Neurons of the lateral vestibular nucleus are inhibited monosynaptically by the Purkinje cells in the cerebellum. • From the vestibular nuclei, secondary axons distribute this information to 4 sites: spinal cord (muscle control), reticular formation (vomiting center), extraocular muscles, and cortex (conscious perception). • Some primary fibers of the vestibular nerve pass directly to the cerebellum, ending in the cortex of the flocculonodular lobe 1. Vestibulospinal Tracts • Both tracts have a strong facilitating effects on motor neurons innervating antigravity muscles • Assists the local myotactic reflexes • Reinforce the tonus of the extensor muscles of the trunk and limbs • Produces enough strength to support the body against gravity and maintain an upright posture • Both terminate along their course almost exclusively upon interneurons in laminae VII and VIII synapse on the alpha and gamma lower motorneurons in Lamina IX • Two major projections into the spinal cord arise from the vestibular nuclei: lateral (maintains upright posture) and medial (coordinated movements of neck & eye) vestibulospinal tract • Lateral vestibular nucleus → uncrossed lateral vestibulospinal tract → cervical to the lumbosacral spinal cord → antigravity ms, neck ms that control gaze • Medial vestibular nucleus → crossed and uncrossed medial vestibulospinal tract → descending portion of MLF (medial longitudinal fasiculus)→ cervical spinal cord only → antigravity ms
Figure 9. Vestibulospinal tract RG, SOPIE, KEIFER, TASIE, FILLE, ELIZ, MAY, JANNA J
V. CENTRAL CONNECTIONS & VESTIBULAR NUCLEI
1. Conjugate Eye Movement: in response to head movement & position of head in space Vestibular nuclei via MLF (crossed and uncrossed) ↓ Nuclei of CNs III, IV, VI: EOMs (conjugate eye movement) and Nucleus of CN XI and anterior horn cells of cervical spinal cord (medial vestibulospinal tract: maintains the position of the head) 2. Maintain upright posture Lateral vestibular nucleus ↓ Lateral vestibulospinal tract of spinal cord to sacral level: extensor muscles of the trunk and limits 3. Fastigial nuclei of the cerebellum 0 0 -‐ Vestibulocerebellar fibers (1 and 2 ) enter cerebellum via juxtarestiform body (a portion of the inferior cerebellar peduncle). Uncrossed efferent fibers from the fastigial nucleus of the cerebellum project to the brainstem also through the juxtarestiform body. Secondary vestibulocerebellar fibers ↓ Fastigial nuclei & cortex of cerebellar vermis, sends efferent fibers to ↓ Lateral vestibular nucleus: facilitating influences on extensor muscle tone via vestibulospinal tract 4. Reticular formation of brainstem o Vomiting center (group of reticular neurons in medulla near dorsal motor nucleus of vagus) § Parasympathetic motor impulses to thoracic & abdominal viscera § Connections from vestibular nuclei to the vomiting center probably account for the vomiting associated with motion sickness o RAS (Reticular Activating System) or Reticular centers § Vestibuloreticular connections to the RAS may alert individual to sleep (rocking chair for babies) 5. MGB (Medial geniculate body) o Vestibular influences project rostrally via the MGB to a vestibular cortical area near the auditory cortex o Related to objective sensations, e.g. dizziness associated with the vestibular system
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A. Test of Vestibular Functions
1.
2.
Rotation test 0 • Subject will sit on a rotating chair with head tilted 30 forward which will then be stopped after 10-‐12 rotations • Induces nystagmus, which lasts about 30 seconds in neurologically normal persons Caloric/Thermal Test • Subject will be tested on both sides 0 • Lying down with head tilted forward about 30 or sitting with 0 head tilted backward 60 to bring the horizontal SCC into a vertical plane • Ear will be irrigated with warm/cold water which will cause a convection current on endolymph which will stimulate the hair cells and cause nystagmus • COWS – direction of nystagmus o Cold on Opposite, warm on Same Side
Reference: 2014A trans Lansang Notes Snell Moore Berne & Levy “Everybody is a genius. But if you judge a fish by its ability to climb a tree, it will live its whole life believing that it is stupid.” -‐-‐ Albert Einstein "If you don't go after what you want, you'll never have it. If you don't ask, the answer is always no. If you don't step forward, you're always in the same place." J
CLINICAL CORRELATIONS A. Nystagmus
Persistent stimulation of hair cells in cristae ampullaris will draw eyes slowly to one side until a limit is reached then jerk quickly to opposite side. (involuntary back and forth, up and down rotational movement of the eyeball) • its direction is designated according to the direction of the fast component which is the basis for tests in vestibular function • this results from a lesion of the vestibular system, its peripheral and central connection and also from lesions in brainstem and cerebellum • this can also be caused by chronic visual impairment or toxic substances • if unbalanced: o Unilateral damage to the vestibular nuclei or their connections → tonic deviation of the eyes to one side + vertigo + nystagmus o Destruction of the vestibular receptors or section of the nerve: (-‐) tonic deviation • Fast case: nystagmus diagnosis o Ex. Lesion on R o Caloric/thermal test: § Cold water-‐direction of nystagmus is opposite the lesion § Warm water-‐direction of nystagmus is to the same side B. Vertigo •
• Sensation of whirling, dizziness due to disturbance of equilibrium • Stimulation or damage to the vestibular end organ (prolonged or excessive stimulation of vestibular apparatus -‐ motion sickness) RG, SOPIE, KEIFER, TASIE, FILLE, ELIZ, MAY, JANNA J
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