Survey
* Your assessment is very important for improving the workof artificial intelligence, which forms the content of this project
* Your assessment is very important for improving the workof artificial intelligence, which forms the content of this project
The Auditory and Vestibular System Chapter 15 The Auditory and Vestibular System Auditory System - sense of hearing Used to detect sound and interpret nuances of sound Important in communication and survival Able to evoke sensations and emotions Vestibular system - sense of balance Informs nervous system about the relative position and movement of the head Subconsciously controls muscle to reorient body and eye position. The Nature of Sound Hearing is a response to vibrating air molecules. Sounds are audible variations in air pressure Moving objects compress air as they move forward and decrease the density of air as they move away. Waves move at 343 m/s or 767 mph. The Nature of Sound Pitch is determined by the frequency of vibration Frequency = the number of compressions per second. One cycle is the distance between the waves of compression Frequency is expressed in hertz (Hz) Hearing range is 20 to 20,000 Hz. Most sensitive to frequencies ranging from 1,500 to 4,000 Hz. Decreases with age or exposure to loud sounds. Loudness = difference in pressure between compressed and rarefied patches of air Range is tremendous loudest sound without ear damage is a trillion times greater than the faintest sound we can hear Intensity is expressed in decibels (dB) 120 to 140 dB causing pain in most people. Real world sounds rarely consist of simple periodic sound waves at one frequency or intensity. The Structure of the Auditory System Outer Ear Auricle or Pinna External Auditory Meatus Tympanic Membrane Middle Ear Ear ossicles Inner Ear Vestibule Cochlea Middle Ear Ear Ossicles Malleus, Incus, Stapes Act as a lever system to amplify sound. Eustachian tubes Equalizes pressure in middle ear Attenuation Reflex Tensor Tympani and Stapedius Muscle Reduces hearing saturation levels Protects the inner ear. Reduces low-frequency background noise. The Inner Ear Vestibule Semicircular Canals Cochlea Chambers Scala Vestibuli Scala Media Scala Tympani Membranes Basilar membrane Reissner’s (vestibular) membrane Tectorial membrane Stria vascularis Cochlear Structures Oval Window Round window Helicotrema Basilar membrane widens toward the Apex of cochlea. Perilymph Fills Scala Vestiblia and Scala Tympani Low K+, High Na+ Endolymph Fills Scala media High K+, Low Na+ Produces an endocochlear potential that enhances auditory transduction Basilar Membrane Structural properties determine how the membrane responds to sound. Wider at apex than at the base by 5 times. Stiffest at base and most flexible at apex. Movement of stapes causes endolymph to flow causing a traveling wave in the membrane The distance the wave travels depends on the frequency of the wave. Different locations of the basilar membrane are maximally deformed at different frequencies creating a placed code. Response of the Basilar Membrane to Sound High frequency waves dissipate near the narrow, stiff base. Low frequency waves dissipate near the wide flexible apex. A place code where maximum amplitude deflection occurs is responsible for the neural coding of pitch. Organ of Corti Hair cells Three rows of outer hair cells and one inner. Stereocilia are embedded in the reticular lamina and tectorial membrane. Have no axons Interact with bipolar spiral ganglion cells that form the cochlear (auditory) nerve. Transduction by Hair Cells. Vibration in the basilar membrane results in the bending of stereocilia Stereocilia are cross linked by filaments and move together as a unit. Bending of steriocilia causes changes in the polarization of the hair cells. Displacement of only 0.3 nm can be detected (diameter of a large molecule). Loud noises that saturate hair cells move stereocilia by only 20 nm. Bending depolarizes or hyperpolarizes hair cells depending on the direction the stereocilia are pulled. Hair cell receptor potentials closely follow the air pressure changes during a lowfrequency sound. Depolarization of Hair Cells. K+ channels on the stereocilia are linked by elastic filaments. Displacement of cilia opens or closes K+ channels K+ entering cell causes depolarization. Note: K+ entry generally causes hyperpolarization. Endolymph has a high concentration of K+. Depolarization causes voltage gated Ca++ channels to open Ca++ triggers the release of neurotransmitter. High K+ Low Na+ High Na+ Low K+ The Innervation of Hair Cells Auditory nerves are bipolar with nuclei in the spiral ganglia. 95% of spiral ganglia neurons communicate with inner hair cells. Each inner hair cells feeds about 10 spiral ganglia cells Most detection of sound occurs on the inner hair cell. One spiral ganglia cell will connect to multiple outer hair cells. Amplification by Outer Hair Cells Motor proteins in the outer hair cells can shorten hair cells. Shortening of hair cells increases the bending of the basilar membrane. Amplification of basilar membrane vibration causes the stereocilia on the inner hair cells to bend more. Central Auditory Processes Spiral ganglion neurons travel through the vestibulo-cochlear nerve to the medulla and branch to enter both the dorsal and ventral cochlear nucleus. Neural signals travel through numerous pathways. The ventral cochlear nucleus projects to the olivary bodies on both sides of the brain then through the lateral lemniscus. The dorsal path bipasses the superior olive. All paths converge at the Inferior Colliculus then go on to the Medial Geniculate Nucleus then into the Auditory Cortex. Central Auditory Processes Inferior colliculus communicates with the superior colliculus to integrate with visual input. There is an extensive feedback system in the auditory system Other than the cochlear nuclei, auditory nuclei receive input from both ears. Response Properties of Spiral Ganglion Cells Spiral ganglion cells are frequency tuned. Each cell responds at a characteristic frequency. Response properties of nuclei are diverse Cochlear nuclei specialized for varying time with frequency MGN- Vocalization Superior Olive Sound localization Encoding Sound Intensity and Frequency Sounds are diverse and complex Our brain must analyze the important ones and ignore the noise Sound is differentiated based upon intensity, frequency and point of origin Each of these features is represented differently in the auditory pathway. Stimulus Frequency and Tonotopy