Audition: Hearing, the Ear, and Sound Localization

The human auditory system allows us to perceive and localize sounds in our physical environment.

The human auditory system allows the body to collect and interpret sound waves into meaningful
messages. The main sensory organ responsible for the ability to hear is the ear, which can be broken
down into the outer ear, middle ear, and inner ear. The inner ear contains the receptor cells necessary
for both hearing and equilibrium maintenance. Human beings also have the special ability of being able
to estimate where sounds originate from, commonly called sound localization.

The Ear
The ear is the main sensory organ of the auditory system. It performs the first processing of sound and
houses all of the sensory receptors required for hearing. The ear's three divisions (outer, middle, and
inner) have specialized functions that combine to allow us to hear.

The outer ear is the external portion of the ear, much of which can be seen on the outside of the human
head. It includes the pinna, the ear canal, and the most superficial layer of the ear drum, the tympanic
membrane. The outer ear's main task is to gather sound energy and amplify sound pressure. The pinna,
the fold of cartilage that surrounds the ear canal, reflects and attenuates sound waves, which helps the
brain determines the location of the sound. The sound waves enter the ear canal, which amplifies the
sound into the ear drum. Once the wave has vibrated the tympanic membrane, sound enters the middle
ear. The middle ear is an air-filled tympanic (drum-like) cavity that transmits acoustic energy from the
ear canal to the cochlea in the inner ear. This is accomplished by a series of three bones in the middle
ear: the malleus, the incus, and the stapes. The malleus (Latin for "hammer") is connected to the mobile
portion of the ear drum. It senses sound vibrations and transfers them onto the incus. The incus (Latin
for "anvil") is the bridge between the malleus and the stapes. The stapes (Latin for "stirrup") transfers
the vibrations from the incus to the oval window, the portion of the inner ear to which it is connected.
Through these steps, the middle ear acts as a gatekeeper to the inner ear, protecting it from damage by
loud sounds. Unlike the middle ear, the inner ear is filled with fluid. When the stapes footplate pushes
down on the oval window in the inner ear, it causes movement in the fluid within the cochlea. The
function of the cochlea is to transform mechanical sound waves into electrical or neural signals for use
in the brain. Within the cochlea there are three fluid-filled spaces: the tympanic canal, the vestibular
canal, and the middle canal. Fluid movement within these canals stimulates hair cells of the organ of
Corti, a ribbon of sensory cells along the cochlea. These hair cells transform the fluid waves into
electrical impulses using cilia, a specialized type of mechanosensor.

The Process of Hearing

Hearing begins with pressure waves hitting the auditory canal and ends when the brain perceives
sounds. Sound reception occurs at the ears, where the pinna collects, reflects, attenuates, or amplifies
sound waves. These waves travel along the auditory canal until they reach the ear drum, which vibrates
in response to the change in pressure caused by the waves. The vibrations of the ear drum cause
oscillations in the three bones in the middle ear, the last of which sets the fluid in the cochlea in motion.
The cochlea separates sounds according to their place on the frequency spectrum. Hair cells in the
cochlea perform the transduction of these sound waves into afferent electrical impulses. Auditory nerve
fibers connected to the hair cells form the spiral ganglion, which transmits the electrical signals along
the auditory nerve and eventually on to the brain stem. The brain responds to these separate
frequencies and composes a complete sound from them.

Sound Localization
Humans are able to hear a wide variety of sound frequencies, from approximately 20 to 20,000 Hz. Our
ability to judge or estimate where a sound originates, called sound localization, is dependent on the
hearing ability of each ear and the exact quality of the sound. Since each ear lies on an opposite side of
the head, a sound reaches the closest ear first, and the sound's amplitude will be larger (and therefore
louder) in that ear. Much of the brain's ability to localize sound depends on these interaural (between-
the-ears) differences in sound intensity and timing. Bushy neurons can resolve time differences as small
as ten milliseconds, or approximately the time it takes for sound to pass one ear and reach the other.

Gustation: Taste Buds and Taste

The gustatory system, including the mouth, tongue, and taste buds, allows us to transduce chemical
molecules into specific taste sensations.

The gustatory system creates the human sense of taste, allowing us to perceive different flavors from
substances that we consume as food and drink. Gustation, along with olfaction (the sense of smell), is
classified as chemoreception because it functions by reacting with molecular chemical compounds in a
given substance. Specialized cells in the gustatory system that are located on the tongue are called taste
buds, and they sense tastants (taste molecules). The taste buds send the information from the tastants
to the brain, where a molecule is processed as a certain taste. There are five main tastes: bitter, salty,
sweet, sour, and umami (savory). All the varieties of flavor we experience are a combination of some or
all of these tastes.

Tongue and Taste Buds

The sense of taste is transduced by taste buds, which are clusters of 50-100 taste receptor cells located
in the tongue, soft palate, epiglottis, pharynx, and esophagus. The tongue is the main sensory organ of
the gustatory system. The tongue contains papillae, or specialized epithelial cells, which have taste buds
on their surface. There are three types of papillae with taste buds in the human gustatory system:
 1. Fungiform papillae, which are mushroom-shaped and located at the tip of the tongue;
2. Foliate papillae, which are ridges and grooves toward the back of the tongue;
3. Circumvallate papillae, which are circular-shaped and located in a row just in front of the end of
the tongue.

Each taste bud is flask-like in shape and formed by two types of cells: supporting cells and gustatory
cells. Gustatory cells are short-lived and are continuously regenerating. They each contain a taste pore
at the surface of the tongue which is the site of sensory transduction. Though there are small
differences in sensation, all taste buds, no matter their location, can respond to all types of taste.

Tastes
Traditionally, humans were thought to have just four main tastes: bitter, salty, sweet, and sour.
Recently, umami, which is the Japanese word for "savory," was added to this list of basic tastes. (Spicy is
not a basic taste because the sensation of spicy foods does not come from taste buds but rather from
heat and pain receptors.) In general, tastes can be appetitive (pleasant) or aversive (unpleasant),
depending on the unique makeup of the material being tasted. There is one type of taste receptor for
each flavor, and each type of taste stimulus is transduced by a different mechanism. Bitter, sweet, and
umami tastes use similar mechanisms based on a G protein-coupled receptor, or GPCR.

 Bitter

There are several classes of bitter compounds which vary in chemical makeup. The human body has
evolved a particularly sophisticated sense for bitter substances and can distinguish between the many
radically different compounds that produce a bitter response. Evolutionary psychologists believe this to
be a result of the role of bitterness in human survival: some bitter-tasting compounds can be hazardous
to our health, so we learned to recognize and avoid bitter substances in general.

 Salty

The salt receptor, NaCl, is arguable the simplest of all the receptors found in the mouth. An ion channel
in the taste cell wall allows Na+ ions to enter the cell. This depolarizes the cell and floods it with ions,
leading to a neurotransmitter release.

 Sweet

Like bitter tastes, sweet taste transduction involves GPCRs binding. The specific mechanism depends on
the specific molecule flavor. Natural sweeteners such as saccharides activate the GPCRs to release
gustducin. Synthetic sweeteners such as saccharin activate a separate set of GPCRs, initiating a similar
but different process of protein transitions.

 Sour

Sour tastes signal the presence of acidic compounds in substances. There are three different receptor
proteins at work in a sour taste. The first is a simple ion channel which allows hydrogen ions to flow
directly into the cell. The second is a K+ channel which has H+ ions in order to block K+ ions from
escaping the cell. The third allows sodium ions to flow down the concentration gradient into the cell.
This involvement with sodium ions implies a relationship between salty and sour tastes receptors.

 Umami

Umami is the newest receptor to be recognized by western scientists in the family of basic tastes. This
Japanese word means "savory" or "meaty." It is thought that umami receptors act similarly to bitter and
sweet receptors (involving GPCRs), but very little is known about their actual function. We do know that
umami detects glutamates that are common in meats, cheese, and other protein-heavy foods and reacts
specifically to foods treated with MSG.

References;

Book; Introduction to Psychology

Morgan T. Clifford
King A. Richard.
Weisz R. John.
Schopler John.
Published by; McGraw Hill Education (India Pvt. Ltd) Edition 1993.

James T. Walker, Arthur L. Irion, Douglas G. Gordon- Attention, Perception &

Psychophysics,1981.