First you tell them what your gonna tell The vestibular system them • The vestibular organs sense head motion: canals sense Michael E. Goldberg, M.D. rotation; otoliths sense linear acceleration (including gravity). • The central vestibular system distributes this signal to oculomotor, head movement, and postural systems for gaze, Please sit where you can examine a partner head, and limb stabilization.. • The visual system complements the vestibular system. • Visuo-vestibular conflict causes acute discomfort. • Peripheral and brainstem vestibular dysfunction causes pathological sense of self-motion and visuo-vestibular conflict. The vestibular labyrinth answers the two The vestibular labyrinth answers two questions basic to the human condition by questions basic to the human condition sensing • Where am I going? • Head angular acceleration (semicircular canals) • Head rotation. • Which way is up? • Head linear acceleration (saccule and utricle) • Translational motion. • Gravity (and by extension head tilt). The vestibular organ lies in the temporal The vestibular organ bone Vestibular Nerve Anterior vertical canal Facial Nerve Horizontal canal Vestibulocochlear (VIII) Nerve Foramen Magnum Posterior vertical canal Cochlear Nerve Cochlear Nerve Cochlea Saccule Utricle 1
Each vestibular organ has a sensor for head Deformation of the stereocilia towards acceleration, driven by hair cells similar to the kinocilium causes hyperpolarization those in the cochlea • In the cochlea vibration induced by sound deforms the hair cells. ⇒ depolarization • In the labyrinth acceleration deforms the hair ⇐ hyperpolarization cells. • In the semicircular canals the sensing organ is the ampulla How the semicircular canals sense Hair cells respond to deformation rotation Hair Cell Cupula Ampullary Crista Ampulla Semicircular Canal Endolymph Vestibular Neuron The three semicircular canals lie in 3 The semicircular canals are functionally orthogonal planes paired and sense rotation • Horizontal canals: rotation in the horizontal Cochlea Cochlear N plane Anterior Vertical • Left anterior and right posterior canals (LARP): Vestibular N Canal rotation in the vertical plane skewed 45° anteriorly to the left. Vestibulo- Horizontal Cochlear N • Right anterior and left posterior canals (RALP): Canal (Nerve VIII) rotation in the vertical plane skewed 45° Posterior anteriorly to the right. Vertical Canal 2
The semicircular canals are functionally The otolith organs sense linear paired acceleration. Hair cells lie in the macula. Otoconia Otolithic • The canals lie in roughly the ��������� �������� membrane same planes as the extraocular Otoconia (ear dust) ��������� �������� muscles: Striola • Horizontal canals: lateral and �������� ��������� �� medial recti. Otolithic �������� ��������� �� �� • LARP: left vertical recti, right Membraine obliques. ����� ���� �������� ��������� �� �� � • RALP: right vertical recti, left ���� ����� ��������� �������� �� � � obliques. • Each canal excites a pair of ���� ����� �������� ��������� �� � � muscles and inhibits a pair of ���� ����� ��������� �������� �� �� muscles in its plane. Its partner When the head tilts the hair cells Hair cells excites the muscles it inhibits, ����� ���� �������� � ��������� � depolarize, are distorted by the shift of the exciting and vice-versa. afferent fibers ����� ��������� � � otolithic membrane Hair cells ����� ��������� hyperpolarize, inhibiting afferent fibers The otolith organs sense linear acceleration • The saccule senses acceleration in the sagittal vertical plane: up and down (so it senses gravity) and forward and backward. Mnemonic: Saccule - Sagittal • The utricle senses acceleration in the horizontal plane: The signals in the vestibular There are 3 major vestibular reflexes nerve • Although the cupula senses • Vestibulo-ocular reflex – keep the eyes still in acceleration, the canal signal in the space when the head moves. vestibular nerve is a tonic signal, • Vestibulo-colic reflex – keeps the head still in deviations from which are proportional space – or on a level plane when you walk. to head velocity. • Vestibular-spinal reflex – adjusts posture for • The macular afferents have a tonic rapid changes in position. signal, deviations from which are sensitive to acceleration. 3
Connections to the vestibular nucleus Nuclear Connections of the Otolith from the canals Organs The Medial Vestibulospinal Tract The lateral vestobulospinal tract (MVST) ● Originates in the lateral vestibular nucleus, predominantly an otolith signal. ● Originates in the medial vestibular ● Projects to cervical, thoracic, and lumbar segmen nucleus, predominantly a canal signal. via the ventral funiculus. ● Predominantly projects to cervical ● Entirely ipsilateral. segments via the medial longitudinal ● Allows the legs to adjust for head movements. fasciculus. ● Provides excitatory tone to extensor muscles. ● Predominantly ipsilateral. ● Decerebrate rigidity is the loss of inhibition from ● Keeps the head still in space – mediating cerebral cortex and cerebellum on the LVST, the vestibulo-colic reflex. and exagerates the effect of the tonic signal in the LVST. The Horizontal Rotational The Horizontal Translational VOR Vestibulo-ocular Reflex • Keeps the eyes still when the head moves laterally Head position (for example when you are looking out of the window of the A train and trying to read the name of the station past which you are traveling). • Gain is dependent on viewing distance: during translation a far object moves less on the retina Eye position than a near object. • The rotational VOR is not dependent upon viewing distance. Gaze position • Most head movement evokes a combination of the rotational (canal) and translation (otolith) VOR’s. 4
The horizontal vestibulo-ocular reflex The VOR is plastic (VOR) • It can be suppressed when you don’t want it. Left Medial Rectus Right Lateral Rectus • Its gain can change. • How do you know if the VOR is doing a good job? Oculomotor • There is no motion on the retina when the head moves. Abducens Nerve (III) Nerve (VI) • If a muscle is weakened, a given central signal will be Oculomotor inadequate, and the world will move on the retina. Nucleus • This can be mimicked by spectacles that increase retinal slip. Abducens • In either case, the brain adjusts the VOR signal so the Vestibular Nuclei Nucleus retinal slip is eliminated. Lateral Medial • The cerebellum is necessary for both suppression of the Nucleus VOR and for slip-induced gain change. Prepositus Hypoglossi The optokinetic signal Vestibular Nystagmus • The vestibular system is imperfect • The cupula habituates in 5 seconds. • The brainstem and cerebellum extend this time to roughly 25 seconds, after which there is no further response to head acceleration. • The vestibular system is a poor transducer of very slow (<0.1Hz) rotation. • The visual system compensates for the inadequacies of the vestibular signal by providing a description of the retinal motion evoked by the head movement. • The optokinetic response is mediated by neurons in the accessory optic system in the pretectum, and the motion- sensitive areas in the cortex (MT and MST). The vestibular nucleus combines Visual-vestibular conflict visual and vestibular signals • Full-field stimulation is an effective stimulus Rotate in Dark for the vestibular nucleus. The neurons can’t tell the difference, nor can you! • Ordinarily the head movement implied by the Rotate in Light visual and visual signals are equal. • Motion sickness – nausea and vomiting – occurs when the visual and vestibular signals Visual Motion are unequal. 5
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