36.4 Hearing and vestibular sensation  (Page 4/28)

Higher processing

The inner hair cells are most important for conveying auditory information to the brain. About 90 percent of the afferent neurons carry information from inner hair cells, with each hair cell synapsing with 10 or so neurons. Outer hair cells connect to only 10 percent of the afferent neurons, and each afferent neuron innervates many hair cells. The afferent, bipolar neurons that convey auditory information travel from the cochlea to the medulla, through the pons and midbrain in the brainstem, finally reaching the primary auditory cortex in the temporal lobe.

Vestibular information

The stimuli associated with the vestibular system are linear acceleration (gravity) and angular acceleration and deceleration. Gravity, acceleration, and deceleration are detected by evaluating the inertia on receptive cells in the vestibular system. Gravity is detected through head position. Angular acceleration and deceleration are expressed through turning or tilting of the head.

The vestibular system has some similarities with the auditory system. It utilizes hair cells just like the auditory system, but it excites them in different ways. There are five vestibular receptor organs in the inner ear: the utricle, the saccule, and three semicircular canals. Together, they make up what’s known as the vestibular labyrinth that is shown in [link] . The utricle and saccule respond to acceleration in a straight line, such as gravity. The roughly 30,000 hair cells in the utricle and 16,000 hair cells in the saccule lie below a gelatinous layer, with their stereocilia projecting into the gelatin. Embedded in this gelatin are calcium carbonate crystals—like tiny rocks. When the head is tilted, the crystals continue to be pulled straight down by gravity, but the new angle of the head causes the gelatin to shift, thereby bending the stereocilia. The bending of the stereocilia stimulates the neurons, and they signal to the brain that the head is tilted, allowing the maintenance of balance. It is the vestibular branch of the vestibulocochlear cranial nerve that deals with balance.

The fluid-filled semicircular canals are tubular loops set at oblique angles. They are arranged in three spatial planes. The base of each canal has a swelling that contains a cluster of hair cells. The hairs project into a gelatinous cap called the cupula and monitor angular acceleration and deceleration from rotation. They would be stimulated by driving your car around a corner, turning your head, or falling forward. One canal lies horizontally, while the other two lie at about 45 degree angles to the horizontal axis, as illustrated in [link] . When the brain processes input from all three canals together, it can detect angular acceleration or deceleration in three dimensions. When the head turns, the fluid in the canals shifts, thereby bending stereocilia and sending signals to the brain. Upon cessation accelerating or decelerating—or just moving—the movement of the fluid within the canals slows or stops. For example, imagine holding a glass of water. When moving forward, water may splash backwards onto the hand, and when motion has stopped, water may splash forward onto the fingers. While in motion, the water settles in the glass and does not splash. Note that the canals are not sensitive to velocity itself, but to changes in velocity, so moving forward at 60mph with your eyes closed would not give the sensation of movement, but suddenly accelerating or braking would stimulate the receptors.

Higher processing

Hair cells from the utricle, saccule, and semicircular canals also communicate through bipolar neurons to the cochlear nucleus in the medulla. Cochlear neurons send descending projections to the spinal cord and ascending projections to the pons, thalamus, and cerebellum. Connections to the cerebellum are important for coordinated movements. There are also projections to the temporal cortex, which account for feelings of dizziness; projections to autonomic nervous system areas in the brainstem, which account for motion sickness; and projections to the primary somatosensory cortex, which monitors subjective measurements of the external world and self-movement. People with lesions in the vestibular area of the somatosensory cortex see vertical objects in the world as being tilted. Finally, the vestibular signals project to certain optic muscles to coordinate eye and head movements.

Section summary

Audition is important for territory defense, predation, predator defense, and communal exchanges. The vestibular system, which is not auditory, detects linear acceleration and angular acceleration and deceleration. Both the auditory system and vestibular system use hair cells as their receptors.

Auditory stimuli are sound waves. The sound wave energy reaches the outer ear (pinna, canal, tympanum), and vibrations of the tympanum send the energy to the middle ear. The middle ear bones shift and the stapes transfers mechanical energy to the oval window of the fluid-filled inner ear cochlea. Once in the cochlea, the energy causes the basilar membrane to flex, thereby bending the stereocilia on receptor hair cells. This activates the receptors, which send their auditory neural signals to the brain.

The vestibular system has five parts that work together to provide the sense of direction, thus helping to maintain balance. The utricle and saccule measure head orientation: their calcium carbonate crystals shift when the head is tilted, thereby activating hair cells. The semicircular canals work similarly, such that when the head is turned, the fluid in the canals bends stereocilia on hair cells. The vestibular hair cells also send signals to the thalamus and to somatosensory cortex, but also to the cerebellum, the structure above the brainstem that plays a large role in timing and coordination of movement.

Art connections