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Hearing and Equilibrium

Author: Sophia

what's covered
In this lesson, you will learn about the special senses of hearing and equilibrium, which are associated with the ear. Specifically, this lesson will cover:

Table of Contents


1. Audition (Hearing)

Audition is the special sense of hearing that makes use of the ear to convert sound waves into neural signals and is associated with the ear.

The large, fleshy structure on the lateral aspect of the head is known as the auricle. The C-shaped curves of the auricle direct sound waves toward the auditory canal. The canal enters the skull through the external auditory meatus of the temporal bone. At the end of the auditory canal is the tympanic membrane, or eardrum, which vibrates after it is struck by sound waves. The auricle, ear canal, and tympanic membrane are often referred to as the external ear.

The middle ear consists of a space spanned by three small bones called the ossicles. The three ossicles are the malleus, incus, and stapes, which are Latin names that roughly translate to hammer, anvil, and stirrup. The malleus is attached to the tympanic membrane and articulates with the incus. The incus, in turn, articulates with the stapes. The stapes is then attached by the oval window (which you will learn more about shortly) to the inner ear, where the sound waves will be transduced into a neural signal.

The middle ear is connected to the pharynx through the auditory tube, also known as the Eustachian tube or pharyngotympanic tube, which helps equilibrate air pressure across the tympanic membrane. The tube is normally closed but will pop open when the muscles of the pharynx contract during swallowing or yawning.

IN CONTEXT

When you travel up or down in elevation, you might notice your ears can be irritated. This irritation is caused by the change in atmospheric pressure.

Under normal conditions, the pressure on both sides of the tympanic membrane (eardrum) are equal, allowing sound waves (i.e., small shifts in pressure) to cause the eardrum to move when they strike it. However, when atmospheric pressure changes by changing elevation or your position in water (sinking to the bottom of a pool or rising up to the surface while scuba diving), the pressure is no longer equal. This can cause irritation and possible damage.

Swallowing or yawning can manipulate the auditory tube connecting the middle ear to the mouth, allowing for air to enter or escape this cavity, and equalizing the pressure across the eardrum.

The inner ear is often described as a bony labyrinth, as it is composed of a series of bony canals embedded within the temporal bone. It has two separate regions, the cochlea and the vestibule, which are responsible for hearing and balance, respectively.

Structures of the Ear—The external ear contains the auricle, ear canal, and tympanic membrane. The middle ear contains the ossicles and is connected to the pharynx by the Eustachian tube. The inner ear contains the cochlea and vestibule, which are responsible for audition and equilibrium, respectively.


You can see that the cochlea looks like a snail's shell, a spiral moving inwards. On the lateral portion of the cochlea is the oval window, an oval-shaped hole in the bone where the stapes sits. Inside the bony labyrinth of the inner ear is a fluid-filled tube that spirals along with the bone to the center. Just inferior to the oval window is the round window, a round-shaped hole in the cochlea covered by a membrane. As sound waves strike the tympanic membrane and move it inwards, the ossicle bones shift. When the stapes moves, it presses into the oval window, causing a fluid wave within the cochlea. The round window will bulge out to accommodate the wave and pucker in when the wave recedes. The frequency of the fluid waves will match the frequency of the sound waves.

Transmission of Sound Waves to the Cochlea—A sound wave causes the tympanic membrane to vibrate. This vibration is amplified as it moves across the malleus, incus, and stapes. The amplified vibration is picked up by the oval window, causing pressure waves in the fluid of the scala vestibuli and scala tympani. The complexity of the pressure waves is determined by the changes in amplitude and frequency of the sound waves entering the ear.


Along the center of the winding cochlea is a cavity called the cochlear duct, which contains an auditory sensory organ called the organ of Corti. As the fluid waves move through the cochlea, the cochlear duct moves at a specific spot, depending on the frequency of the waves. Higher frequency waves move the region of the duct that is close to the base of the cochlea. Lower frequency waves move the region that is near the tip of the cochlea.


The organ of Corti contains sensory receptor cells that are specialized mechanoreceptors called hair cells, which are named for the hair-like stereocilia extending from the cell’s apical surfaces. The stereocilia are an array of microvilli-like structures arranged from tallest to shortest. Protein fibers tether adjacent hairs together within each array, such that the array will bend in response to movements of the underlying tissue.

The stereocilia extend up from the hair cells to the overlying tectorial membrane, which is attached medially to the organ of Corti. When the pressure waves move the organ of Corti, the tectorial membrane slides across the stereocilia. This bends the stereocilia either toward or away from the tallest member of each array. When the stereocilia bend toward the tallest member of their array, tension in the protein tethers opens ion channels in the hair cell membrane. This will depolarize the hair cell membrane. When the stereocilia bend toward the shortest member of their array, the tension on the tethers slackens and the ion channels close.

When no sound is present, and the stereocilia are standing straight, a small amount of tension still exists on the tethers, keeping the membrane potential of the hair cell slightly depolarized. Depolarized hair cells send signals to the auditory sensory neuron. All auditory sensory neurons leave the cochlea as part of the cochlear nerve, which is the auditory portion of the vestibulocochlear nerve (CN VIII).

Hair Cell—The hair cell is a mechanoreceptor with an array of stereocilia emerging from its apical surface. The stereocilia are tethered together by proteins that open ion channels when the array is bent toward the tallest member of their array and closed when the array is bent toward the shortest member of their array.


As stated above, a given region of the cochlear duct will only move if the incoming sound is at a specific frequency. Because the tectorial membrane only moves where the cochlear duct moves, the hair cells in this region will also only respond to sounds of this specific frequency. Therefore, as the frequency of a sound changes, different hair cells are activated all along the basilar membrane.

did you know
The cochlea encodes auditory stimuli for frequencies between 20 and 20,000 Hz, which is the range of sound that human ears can detect.

The unit of Hertz measures the frequency of sound waves in terms of cycles produced per second. Frequencies as low as 20 Hz are detected by hair cells at the apex, or tip, of the cochlea. Frequencies in the higher ranges of 20 KHz are encoded by hair cells at the base of the cochlea, close to the round and oval windows. Most auditory stimuli contain a mixture of sounds at a variety of frequencies and intensities (represented by the amplitude of the sound wave). The hair cells along the length of the cochlear duct, which are each sensitive to a particular frequency, allow the cochlea to separate auditory stimuli by frequency just as a prism separates visible light into its component colors.

Frequency Coding in the Cochlea—The standing sound wave generated in the cochlea by the movement of the oval window deflects the basilar membrane on the basis of the frequency of sound. Therefore, hair cells at the base of the cochlea are activated only by high frequencies, whereas those at the apex of the cochlea are activated only by low frequencies.


terms to know
Audition
The special sense of hearing.
Auricle
The fleshy portion of the external ear.
Tympanic Membrane
The membrane at the end of the external acoustic meatus; eardrum.
External Ear
A portion of the auditory anatomy including the auricle, ear canal, and tympanic membrane.
Middle Ear
A portion of the auditory anatomy between the tympanic membrane and bony labyrinth, including the ossicles.
Ossicles
Three small bones located in the middle ear.
Malleus
The ossicle attached to the tympanic membrane; the hammer.
Incus
The ossicle that connects the malleus and stapes; the anvil.
Stapes
The ossicle that connects to the bony labyrinth of the inner ear; the stirrup.
Inner Ear
The portion of the auditory and vestibular anatomy including the cochlea and vestibule.
Cochlea
The auditory portion of the inner ear.
Vestibule
The vestibular portion of the inner ear.
Oval Window
A membrane-covered hole in the cochlea where the stapes is attached.
Round Window
A membrane-covered hole in the cochlea inferior to the oval window.
Cochlear Duct
A cavity within the cochlea that contains the organ of Corti.
Organ of Corti
A structure within the cochlear duct that contains hair cells and can send auditory sensory signals.
Hair Cell
Sensory receptor cells of the inner ear that detect changes in the position of their stereocilia.
Stereocilia
Microvilli-like extensions of the hair cell membrane that attach to the tectorial membrane.
Tectorial Membrane
A structure of the organ of Corti that stereocilia are connected to.
Auditory Sensory Neuron
Afferent sensory neurons in the ear that are sensitive to hair cell signals; part of the cochlear nerve.
Cochlear Nerve
The auditory branch of the vestibulocochlear nerve.


2. Vestibulation (Equilibrium, Balance)

Along with audition, the inner ear is responsible for encoding information about vestibulation, also known as the sense of equilibrium or balance. Recall that the inner ear is a bony labyrinth composed of the cochlea and vestibule. The cochlea contains auditory structures, whereas the vestibule contains vestibular structures. Within the vestibule are three semicircular canals that detect rotational movement as well as the utricle and saccule that detect linear movement and position.


The utricle and saccule both contain hair cells (which are mechanoreceptors) embedded in them. The stereocilia of the hair cells extend into a viscous gel called the otolithic membrane. On top of the otolithic membrane is a layer of calcium carbonate crystals called otoliths, also known as ear stones.

The otoliths essentially make the otolithic membrane top-heavy. The otolithic membrane moves separately from the underlying tissue in response to head movements. Tilting the head causes the otolithic membrane to slide in the direction of gravity. The moving otolithic membrane, in turn, bends the stereocilia, causing some hair cells to depolarize as others hyperpolarize. The exact position of the head is interpreted by the brain based on the pattern of hair-cell depolarization.

The utricle and saccule can also detect changes in linear movement. The utricle is positioned in the horizontal plane and can detect horizontal acceleration or deceleration such as your car picking up speed or slowing down. The saccule is positioned vertically and can detect vertical acceleration or deceleration such as an elevator.

Linear Acceleration Coding—The utricle and saccule are specialized for sensing linear acceleration, such as when gravity acts on the tilting head or if the head starts moving in a straight line. The difference in inertia between the hair cell stereocilia and the otolithic membrane in which they are embedded leads to a shearing force that causes the stereocilia to bend in the direction of that linear acceleration.


The semicircular canals are three ring-like extensions of the vestibule. One is oriented in the horizontal plane, whereas the other two are oriented in the vertical plane. The base of each semicircular canal, where it meets with the vestibule, connects to an enlarged region known as the ampulla (of the inner ear), which contains hair cells. These hair cells have stereocilia embedded into the cupula, a membrane that attaches to the top of the ampulla.


When the head moves in a rotational movement (i.e., spinning in circles, doing a somersault, or doing a cartwheel), fluid inside one or more of the semicircular canals shifts. As this fluid moves along the ampulla, it bends the cupula and in turn, the stereocilia. This depolarizes the hair cells and sends a signal to a connected ampullary nerve, which extends as the vestibular nerve. This nerve combines with the cochlear nerve to become the vestibulocochlear nerve.

By comparing the signals from all ampullae, the vestibular system can detect the direction of most head movements within three-dimensional (3-D) space.

Rotational Coding by Semicircular Canals—Rotational movement of the head is encoded by the hair cells in the base of the semicircular canals. As one of the canals moves in an arc with the head, the internal fluid moves in the opposite direction, causing the cupula and stereocilia to bend. The movement of two canals within a plane results in information about the direction in which the head is moving, and activation of all six canals can give a very precise indication of head movement in three dimensions.


terms to know
Vestibulation
The sense of equilibrium or balance.
Utricle
A structure of the inner ear able to detect head position and horizontal acceleration and deceleration.
Saccule
A structure of the inner ear able to detect head position and vertical acceleration and deceleration.
Otolithic Membrane
A viscous gelatinous substance in the utricle and saccule of the inner ear that stereocilia of hair cells are embedded in.
Otolith
Calcium carbonate crystals located on top of the otolithic membrane.
Semicircular Canal
Structures of the inner ear that detect rotational movement of the head.
Ampulla (of the Inner Ear)
The enlarged region at the base of a semicircular canal containing hair cells.
Cupula
A membrane located in the ampulla of the semicircular canals where the stereocilia of hair cells are embedded.
Vestibular Nerve
The vestibular branch of the vestibulocochlear nerve.

Interactive 3-D Model
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summary
In this lesson, you learned about the special senses of audition and vestibulation, which are associated with the ear. You first examined the structure of the ear and how it detects sound waves to produce audition (hearing). Then, you learned about how vestibulation (equilibrium, balance) occurs by the structures of the ear detecting body position and body movement.

SOURCE: THIS TUTORIAL HAS BEEN ADAPTED FROM OPENSTAX “ANATOMY AND PHYSIOLOGY 2E”. ACCESS FOR FREE AT OPENSTAX.ORG/BOOKS/ANATOMY-AND-PHYSIOLOGY-2E/PAGES/1-INTRODUCTION. LICENSE: CREATIVE COMMONS ATTRIBUTION 4.0 INTERNATIONAL.

Terms to Know
Ampulla (of the Inner Ear)

The enlarged region at the base of a semicircular canal containing hair cells.

Audition

The special sense of hearing.

Auditory Sensory Neuron

Afferent sensory neurons in the ear that are sensitive to hair cell signals; part of the cochlear nerve.

Auricle

The fleshy portion of the external ear.

Cochlea

The auditory portion of the inner ear.

Cochlear Duct

A cavity within the cochlea that contains the organ of Corti.

Cochlear Nerve

The auditory branch of the vestibulocochlear nerve.

Cupula

A membrane located in the ampulla of the semicircular canals where the stereocilia of hair cells are embedded.

External Ear

A portion of the auditory anatomy including the auricle, ear canal, and tympanic membrane.

Hair Cell

Sensory receptor cells of the inner ear that detect changes in the position of their stereocilia.

Incus

The ossicle that connects the malleus and stapes; the anvil.

Inner Ear

The portion of the auditory and vestibular anatomy including the cochlea and vestibule.

Malleus

The ossicle attached to the tympanic membrane; the hammer.

Middle Ear

A portion of the auditory anatomy between the tympanic membrane and bony labyrinth, including the ossicles.

Organ of Corti

A structure within the cochlear duct that contains hair cells and can send auditory sensory signals.

Ossicles

Three small bones located in the middle ear.

Otolith

Calcium carbonate crystals located on top of the otolithic membrane.

Otolithic Membrane

A viscous gelatinous substance in the utricle and saccule of the inner ear that stereocilia of hair cells are embedded in. 

Oval Window

A membrane-covered hole in the cochlea where the stapes is attached.

Round Window

A membrane-covered hole in the cochlea inferior to the oval window.

Saccule

A structure of the inner ear able to detect head position and vertical acceleration and deceleration.

Semicircular Canal

Structures of the inner ear that detect rotational movement of the head.

Stapes

The ossicle that connects to the bony labyrinth of the inner ear; the stirrup.

Stereocilia

Microvilli-like extensions of the hair cell membrane that attach to the tectorial membrane.

Tectorial Membrane

A structure of the organ of Corti that stereocilia are connected to.

Tympanic Membrane

The membrane at the end of the external acoustic meatus; eardrum.

Utricle

A structure of the inner ear able to detect head position and horizontal acceleration and deceleration.

Vestibular Nerve

The vestibular branch of the vestibulocochlear nerve.

Vestibulation

The sense of equilibrium or balance.

Vestibule

The vestibular portion of the inner ear.