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Sense Organs / 243

               right angles to one another and are desig­  would continually slip across the retina
                                                        during head movement, and focusing on
               nated  anterior,  posterior, and  lateral  to
  VetBooks.ir  describe their orientation. As an extension   the visual field would be difficult or
                                                        impossible while  the  head  was  moving.
               of the membranous labyrinth, each duct is
               filled with endolymph and surrounded by   The vestibular nuclei use information about
               perilymph.                               acceleration to coordinate extraocular
                  One end of each duct is dilated to form   muscle movements with movements of the
               an ampulla, within which are housed the   head and thereby fix the visual image in
               receptor organs of the semicircular ducts   one place on the retina. When the excursion
               (Fig. 12‐10). One wall of the ampulla  features   of the moving head carries the fixed image
               a transverse ridge of connective tissue, the   out of visual range, the eyes dart ahead in
               crista ampullaris, which supports a neu­  the direction of movement to fix upon a
               roepithelium of hair cells. Attached to the   new image. This new image is held on the
               crista is a gelatinous cupula; this extends   retina while the head continues turning
               across the ampulla, forming a flexible   until a compensatory jump ahead is again
               barrier to the flow of endolymph. The cilia   needed. This mechanism results in a cycle
               of the hair cells are embedded in the cupula   of slow movement opposite the direction
               and are therefore bent by movements of it.  of turn (eyes fixed on target) followed by a
                  The semicircular ducts detect angular   rapid readjustment in the same direction
               acceleration (rotation), and their planes   as the turn. This oscillatory reflex eye
               of orientation roughly correspond to the   movement is called nystagmus.
               X‐, Y‐, and Z‐axes of three‐dimensional     Nystagmus is a normal reflex, generated
               space. When the head rotates, the semicir­  in response to movement of the head.
               cular duct lying in that rotational plane   Nystagmus is considered abnormal when it
               moves with it. The endolymph inside the   occurs in absence of head movement; this
               duct, however, must overcome its inertia   resting (spontaneous) nystagmus is a sign
               at the start of rotation and, as a conse­  of vestibular disease.
               quence, briefly lags behind the movement    Axons of some neurons in the vestibular
               of the head. The flexible cupula, acting   nuclei project caudad in a motor tract that
               as  a  dam  across  the  ampulla,  bulges  in   influences activity in  cervical  and  upper
               response to the endolymph’s push, and in   thoracic spinal cord segments. These motor
               doing so, bends the cilia of the embedded   connections activate neck musculature
               hair cells. With three pairs (right and left)   and forelimb extensors, producing the
               of semicircular ducts detecting movement   vestibulocollic reflex,  which  generates
               in the three planes of space, complex rota­  neck movements and forelimb extension to
               tional movements of the head are encoded   help keep the head level with respect to
               in the firing patterns of the six cristae   gravity and movement.
               ampullares.                                 Some fibers from the vestibular nuclei
                  Primary afferent neurons synapse with   form an ipsilateral descending motor tract
               the hair cells of the vestibular apparatus.   that extends the length of the spinal cord.
               Their cell bodies are in the vestibular gan­  This lateral vestibulospinal tract is part
               glion, and their axons constitute the ves­  of the ventromedial motor system (see
               tibular nerve, which joins the cochlear   Chapter  10). Its activity has the primary
               nerve to become the eighth cranial nerve.   effect of increasing tone in antigravity
               Most axons in the vestibular nerve synapse   muscles (proximal limb extensors and axial
               in the large vestibular nuclei of the pons   muscles). This vestibulospinal reflex uses
               and rostral medulla.                     vestibular information to produce limb
                                                        and trunk movements that counteract the
               Vestibular Reflexes.  If  there  were  no   displacement of the head that elicits it. This
               mechanism to keep the eyes steady on a   mechanism is designed to prevent tilting or
               target when the head moved, visual images   falling with shifts in head position.
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