Audition (Hearing)

Hearing, or audition, is the transduction of sound waves into a neural signal that is made possible by the structures of the ear (Figure 15.3.1). The large, fleshy structure on the lateral aspect of the head is known as the auricle or pinna. 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 ear drum, 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.

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 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 Eustachian 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.

The inner ear is composed of a series of canals embedded within the temporal bone often referred to as the bony labyrinth. It has two separate regions, the cochlea and the vestibule, which are responsible for hearing and balance, respectively. These two distinct bundles travel together from the inner ear to the brain stem as the vestibulocochlear nerve.

Sound is transduced into neural signals within the cochlear region of the inner ear, which contains the sensory neurons located within the spiral-shaped cochlea of the inner ear. The cochlea is attached to the stapes through the oval window.

The oval window is located at the beginning of a fluid-filled tube within the cochlea called the scala vestibuli. The scala vestibuli extends from the oval window, travelling above the cochlear duct, which is the central cavity of the cochlea that contains the sound-transducing neurons.

At the uppermost tip of the cochlea, the scala vestibuli curves over the top of the cochlear duct. The fluid-filled tube, now called the scala tympani, returns to the base of the cochlea, this time travelling under the cochlear duct. The scala tympani ends at the round window, which is covered by a membrane that contains the fluid within the scala. As vibrations of the ossicles travel through the oval window, the fluid of the scala vestibuli and scala tympani moves in a wave-like motion. The frequency of the fluid waves match the frequencies of the sound waves (Figure 15.3.2). The membrane covering the round window will bulge out or pucker in with the movement of the fluid within the scala tympani.

A cross-sectional view of the cochlea shows that the scala vestibuli and scala tympani run along both sides of the cochlear duct (Figure 15.3.3). The cochlear duct contains several organs of Corti, which transduce the wave motion of the two scala into neural signals.

The organs of Corti contain hair cells, (stereocilia). (Figure 15.3.4). The stereocilia are microvilli-like structures arranged from tallest to shortest. Protein fibers attach hairs together within each array, such that the array will bend in response to movements of the basilar membrane.

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 from the scala move the basilar membrane, the tectorial membrane slides across the stereocilia. This bends the stereocilia either toward or away from the tallest member of each array.

A given region of the basilar membrane will only move if the incoming sound is at a specific frequency. Because the tectorial membrane only moves where the basilar membrane 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. 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.

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 colours.

 Central Processing of Hearing Information

The sensory pathway for audition travels along the vestibulocochlear nerve, which synapses with neurons in the cochlear nuclei of the superior medulla. Within the brain stem, input from either ear is combined to extract location information from the auditory stimuli. Whereas the initial auditory stimuli received at the cochlea strictly represent the frequency—or pitch—of the stimuli, the locations of sounds can be determined by comparing information arriving at both ears.

Auditory processing continues on to a nucleus in the midbrain called the inferior colliculus. Axons from the inferior colliculus project to two locations, the thalamus and the superior colliculus. The medial geniculate nucleus of the thalamus receives the auditory information and then projects that information to the auditory cortex in the temporal lobe of the cerebral cortex. The superior colliculus receives input from the visual and somatosensory systems, as well as the ears, to initiate stimulation of the muscles that turn the head and neck toward the auditory stimulus.