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Hearing, or auditory perception, is the ability to perceive
sound In physics, sound is a vibration that propagates as an acoustic wave through a transmission medium such as a gas, liquid or solid. In human physiology and psychology, sound is the ''reception'' of such waves and their ''perception'' by the br ...
s through an organ, such as an ear, by detecting
vibration Vibration () is a mechanical phenomenon whereby oscillations occur about an equilibrium point. Vibration may be deterministic if the oscillations can be characterised precisely (e.g. the periodic motion of a pendulum), or random if the os ...
s as periodic changes in the pressure of a surrounding medium. The academic field concerned with hearing is auditory science. Sound may be heard through
solid Solid is a state of matter where molecules are closely packed and can not slide past each other. Solids resist compression, expansion, or external forces that would alter its shape, with the degree to which they are resisted dependent upon the ...
,
liquid Liquid is a state of matter with a definite volume but no fixed shape. Liquids adapt to the shape of their container and are nearly incompressible, maintaining their volume even under pressure. The density of a liquid is usually close to th ...
, or gaseous matter. It is one of the traditional five
sense A sense is a biological system used by an organism for sensation, the process of gathering information about the surroundings through the detection of Stimulus (physiology), stimuli. Although, in some cultures, five human senses were traditio ...
s. Partial or total inability to hear is called hearing loss. In humans and other vertebrates, hearing is performed primarily by the
auditory system The auditory system is the sensory system for the sense of hearing. It includes both the ear, sensory organs (the ears) and the auditory parts of the sensory system. System overview The outer ear funnels sound vibrations to the eardrum, incre ...
: mechanical waves, known as vibrations, are detected by the ear and transduced into nerve impulses that are perceived by the
brain The brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It consists of nervous tissue and is typically located in the head (cephalization), usually near organs for ...
(primarily in the
temporal lobe The temporal lobe is one of the four major lobes of the cerebral cortex in the brain of mammals. The temporal lobe is located beneath the lateral fissure on both cerebral hemispheres of the mammalian brain. The temporal lobe is involved in pr ...
). Like
touch The somatosensory system, or somatic sensory system is a subset of the sensory nervous system. The main functions of the somatosensory system are the perception of external stimuli, the perception of internal stimuli, and the regulation of bo ...
, audition requires sensitivity to the movement of molecules in the world outside the organism. Both hearing and touch are types of mechanosensation.


Hearing mechanism

There are three main components of the human
auditory system The auditory system is the sensory system for the sense of hearing. It includes both the ear, sensory organs (the ears) and the auditory parts of the sensory system. System overview The outer ear funnels sound vibrations to the eardrum, incre ...
: the outer ear, the middle ear, and the inner ear.


Outer ear

The outer ear includes the pinna, the visible part of the ear, as well as the
ear canal The ear canal (external acoustic meatus, external auditory meatus, EAM) is a pathway running from the outer ear to the middle ear. The adult human ear canal extends from the auricle to the eardrum and is about in length and in diameter. S ...
, which terminates at the eardrum, also called the tympanic membrane. The pinna serves to focus sound waves through the ear canal toward the eardrum. Because of the asymmetrical character of the outer ear of most mammals, sound is filtered differently on its way into the ear depending on the location of its origin. This gives these animals the ability to localize sound vertically. The eardrum is an airtight membrane, and when sound waves arrive there, they cause it to vibrate following the
waveform In electronics, acoustics, and related fields, the waveform of a signal is the shape of its Graph of a function, graph as a function of time, independent of its time and Magnitude (mathematics), magnitude Scale (ratio), scales and of any dis ...
of the sound. Cerumen (ear wax) is produced by ceruminous and
sebaceous gland A sebaceous gland or oil gland is a microscopic exocrine gland in the skin that opens into a hair follicle to secrete an oily or waxy matter, called sebum, which lubricates the hair and skin of mammals. In humans, sebaceous glands occur in ...
s in the skin of the human ear canal, protecting the ear canal and tympanic membrane from physical damage and microbial invasion.


Middle ear

The middle ear consists of a small air-filled chamber that is located medial to the eardrum. Within this chamber are the three smallest bones in the body, known collectively as the ossicles which include the malleus, incus, and stapes (also known as the hammer, anvil, and stirrup, respectively). They aid in the transmission of the vibrations from the eardrum into the inner ear, the
cochlea The cochlea is the part of the inner ear involved in hearing. It is a spiral-shaped cavity in the bony labyrinth, in humans making 2.75 turns around its axis, the modiolus (cochlea), modiolus. A core component of the cochlea is the organ of Cort ...
. The purpose of the middle ear ossicles is to overcome the impedance mismatch between air waves and cochlear waves, by providing
impedance matching In electrical engineering, impedance matching is the practice of designing or adjusting the input impedance or output impedance of an electrical device for a desired value. Often, the desired value is selected to maximize power transfer or ...
. Also located in the middle ear are the stapedius muscle and tensor tympani muscle, which protect the hearing mechanism through a stiffening reflex. The stapes transmits sound waves to the inner ear through the oval window, a flexible membrane separating the air-filled middle ear from the fluid-filled inner ear. The round window, another flexible membrane, allows for the smooth displacement of the inner ear fluid caused by the entering sound waves.


Inner ear

The inner ear consists of the
cochlea The cochlea is the part of the inner ear involved in hearing. It is a spiral-shaped cavity in the bony labyrinth, in humans making 2.75 turns around its axis, the modiolus (cochlea), modiolus. A core component of the cochlea is the organ of Cort ...
, which is a spiral-shaped, fluid-filled tube. It is divided lengthwise by the organ of Corti, which is the main organ of mechanical to neural transduction. Inside the organ of Corti is the basilar membrane, a structure that vibrates when waves from the middle ear propagate through the cochlear fluid – endolymph. The basilar membrane is tonotopic, so that each frequency has a characteristic place of resonance along it. Characteristic frequencies are high at the basal entrance to the cochlea, and low at the apex. Basilar membrane motion causes
depolarization In biology, depolarization or hypopolarization is a change within a cell (biology), cell, during which the cell undergoes a shift in electric charge distribution, resulting in less negative charge inside the cell compared to the outside. Depolar ...
of the hair cells, specialized auditory receptors located within the organ of Corti. While the hair cells do not produce
action potential An action potential (also known as a nerve impulse or "spike" when in a neuron) is a series of quick changes in voltage across a cell membrane. An action potential occurs when the membrane potential of a specific Cell (biology), cell rapidly ri ...
s themselves, they release neurotransmitter at synapses with the fibers of the auditory nerve, which does produce action potentials. In this way, the patterns of oscillations on the basilar membrane are converted to spatiotemporal patterns of firings which transmit information about the sound to the
brainstem The brainstem (or brain stem) is the posterior stalk-like part of the brain that connects the cerebrum with the spinal cord. In the human brain the brainstem is composed of the midbrain, the pons, and the medulla oblongata. The midbrain is conti ...
.


Neuronal

The sound information from the cochlea travels via the auditory nerve to the cochlear nucleus in the
brainstem The brainstem (or brain stem) is the posterior stalk-like part of the brain that connects the cerebrum with the spinal cord. In the human brain the brainstem is composed of the midbrain, the pons, and the medulla oblongata. The midbrain is conti ...
. From there, the signals are projected to the
inferior colliculus The inferior colliculus (IC) (Latin for ''lower hill'') is the principal midbrain nucleus of the Auditory system, auditory pathway and receives input from several peripheral brainstem nuclei in the auditory pathway, as well as inputs from the aud ...
in the midbrain tectum. The
inferior colliculus The inferior colliculus (IC) (Latin for ''lower hill'') is the principal midbrain nucleus of the Auditory system, auditory pathway and receives input from several peripheral brainstem nuclei in the auditory pathway, as well as inputs from the aud ...
integrates auditory input with limited input from other parts of the brain and is involved in subconscious reflexes such as the auditory
startle response In animals, including humans, the startle response is a largely unconscious defensive response to sudden or threatening Stimulus (physiology), stimuli, such as sudden noise or sharp movement, and is associated with negative Affect (psychology), af ...
. The inferior colliculus in turn projects to the medial geniculate nucleus, a part of the
thalamus The thalamus (: thalami; from Greek language, Greek Wikt:θάλαμος, θάλαμος, "chamber") is a large mass of gray matter on the lateral wall of the third ventricle forming the wikt:dorsal, dorsal part of the diencephalon (a division of ...
where sound information is relayed to the primary auditory cortex in the
temporal lobe The temporal lobe is one of the four major lobes of the cerebral cortex in the brain of mammals. The temporal lobe is located beneath the lateral fissure on both cerebral hemispheres of the mammalian brain. The temporal lobe is involved in pr ...
. Sound is believed to first become consciously experienced at the primary auditory cortex. Around the primary auditory cortex lies Wernickes area, a cortical area involved in interpreting sounds that is necessary to understand spoken words. Disturbances (such as
stroke Stroke is a medical condition in which poor cerebral circulation, blood flow to a part of the brain causes cell death. There are two main types of stroke: brain ischemia, ischemic, due to lack of blood flow, and intracranial hemorrhage, hemor ...
or trauma) at any of these levels can cause hearing problems, especially if the disturbance is bilateral. In some instances it can also lead to auditory hallucinations or more complex difficulties in perceiving sound.


Hearing tests

Hearing can be measured by behavioral tests using an audiometer. Electrophysiological tests of hearing can provide accurate measurements of hearing thresholds even in unconscious subjects. Such tests include auditory brainstem evoked potentials (ABR), otoacoustic emissions (OAE) and electrocochleography (ECochG). Technical advances in these tests have allowed hearing screening for infants to become widespread. Hearing can be measured by mobile applications which includes audiological hearing test function or hearing aid application. These applications allow the user to measure hearing thresholds at different frequencies ( audiogram). Despite possible errors in measurements, hearing loss can be detected.


Hearing loss

There are several different types of hearing loss: conductive hearing loss,
sensorineural hearing loss Sensorineural hearing loss (SNHL) is a type of hearing loss in which the root cause lies in the inner ear, sensory organ (cochlea and associated structures), or the vestibulocochlear nerve (Cranial nerves, cranial nerve VIII). SNHL accounts for a ...
and mixed types. Recently, the term of Aural Diversity has come into greater use, to communicate hearing loss and differences in a less negatively-associated term. There are defined degrees of hearing loss: * Mild hearing loss - People with mild hearing loss have difficulties keeping up with conversations, especially in noisy surroundings. The most quiet sounds that people with mild hearing loss can hear with their better ear are between 25 and 40 dB HL. * Moderate hearing loss - People with moderate hearing loss have difficulty keeping up with conversations when they are not using a hearing aid. On average, the most quiet sounds heard by people with moderate hearing loss with their better ear are between 40 and 70 dB HL. * Severe hearing loss - People with severe hearing loss depend on powerful hearing aid. However, they often rely on lip-reading even when they are using hearing aids. The most quiet sounds heard by people with severe hearing loss with their better ear are between 70 and 95 dB HL. * Profound hearing loss - People with profound hearing loss are very hard of hearing and they mostly rely on lip-reading and sign language. The most quiet sounds heard by people with profound hearing loss with their better ear are from 95 dB HL or more.


Causes

*Heredity *Congenital conditions *Presbycusis *Acquired **Noise-induced hearing loss **Ototoxic drugs and chemicals **Infection


Prevention

Hearing protection is the use of devices designed to prevent noise-induced hearing loss (NIHL), a type of post-lingual hearing impairment. The various means used to prevent hearing loss generally focus on reducing the levels of noise to which people are exposed. One way this is done is through environmental modifications such as acoustic quieting, which may be achieved with as basic a measure as lining a room with curtains, or as complex a measure as employing an
anechoic chamber An anechoic chamber (''an-echoic'' meaning "non-reflective" or "without echoes") is a room designed to stop reflection (physics), reflections or Echo (phenomenon), echoes of either sound or electromagnetic waves. They are also often isolate ...
, which absorbs nearly all sound. Another means is the use of devices such as
earplug An earplug is a device that is inserted in the ear canal to protect the user's ears from loud noises, intrusion of water, foreign bodies, dust or excessive wind. Earplugs may be used as well to improve sleep quality or focus in noisy environments ...
s, which are inserted into the ear canal to block noise, or
earmuffs Earmuffs are accessories designed to cover a person's ears either for warmth or for Hearing protection device, hearing protection. Both types of earmuff consist of a plastic, thermoplastic or metal head-band that fits over the top or back of the ...
, objects designed to cover a person's ears entirely.


Management

The loss of hearing, when it is caused by neural loss, cannot presently be cured. Instead, its effects can be mitigated by the use of audioprosthetic devices, i.e. hearing assistive devices such as hearing aids and cochlear implants. In a clinical setting, this management is offered by otologists and audiologists.


Relation to health

Hearing loss is associated with
Alzheimer's disease Alzheimer's disease (AD) is a neurodegenerative disease and the cause of 60–70% of cases of dementia. The most common early symptom is difficulty in remembering recent events. As the disease advances, symptoms can include problems wit ...
and
dementia Dementia is a syndrome associated with many neurodegenerative diseases, characterized by a general decline in cognitive abilities that affects a person's ability to perform activities of daily living, everyday activities. This typically invo ...
with a greater degree of hearing loss tied to a higher risk. There is also an association between type 2 diabetes and hearing loss.


Hearing underwater

Hearing threshold and the ability to localize sound sources are reduced underwater in humans, but not in aquatic animals, including whales, seals, and fish which have ears adapted to process water-borne sound.


In vertebrates

Not all sounds are normally audible to all animals. Each species has a range of normal hearing for both amplitude and
frequency Frequency is the number of occurrences of a repeating event per unit of time. Frequency is an important parameter used in science and engineering to specify the rate of oscillatory and vibratory phenomena, such as mechanical vibrations, audio ...
. Many animals use sound to communicate with each other, and hearing in these species is particularly important for survival and reproduction. In species that use sound as a primary means of communication, hearing is typically most acute for the range of pitches produced in calls and speech.


Frequency range

Frequencies capable of being heard by humans are called audio or sonic. The range is typically considered to be between 20 Hz and 20,000 Hz. Frequencies higher than audio are referred to as ultrasonic, while frequencies below audio are referred to as infrasonic. Some
bats Bats are flying mammals of the order Chiroptera (). With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most birds, flying with their very long spread-out ...
use ultrasound for echolocation while in flight. Dogs are able to hear ultrasound, which is the principle of 'silent' dog whistles.
Snake Snakes are elongated limbless reptiles of the suborder Serpentes (). Cladistically squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales much like other members of the group. Many species of snakes have s ...
s sense infrasound through their jaws, and baleen
whale Whales are a widely distributed and diverse group of fully Aquatic animal, aquatic placental mammal, placental marine mammals. As an informal and Colloquialism, colloquial grouping, they correspond to large members of the infraorder Cetacea ...
s, giraffes,
dolphin A dolphin is an aquatic mammal in the cetacean clade Odontoceti (toothed whale). Dolphins belong to the families Delphinidae (the oceanic dolphins), Platanistidae (the Indian river dolphins), Iniidae (the New World river dolphins), Pontopori ...
s and
elephant Elephants are the largest living land animals. Three living species are currently recognised: the African bush elephant ('' Loxodonta africana''), the African forest elephant (''L. cyclotis''), and the Asian elephant ('' Elephas maximus ...
s use it for communication. Some
fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
have the ability to hear more sensitively due to a well-developed, bony connection between the ear and their swim bladder. This "aid to the deaf" for fishes appears in some species such as
carp The term carp (: carp) is a generic common name for numerous species of freshwater fish from the family (biology), family Cyprinidae, a very large clade of ray-finned fish mostly native to Eurasia. While carp are prized game fish, quarries and a ...
and herring.


Time discrimination

Human perception of audio signal time separation has been measured to less than 10 microseconds (10μs). This does not mean that frequencies above are audible, but that time discrimination is not directly coupled with frequency range. Georg Von Békésy in 1929 identifying sound source directions suggested humans can resolve timing differences of 10μs or less. In 1976 Jan Nordmark's research indicated inter-aural resolution better than 2μs. Milind Kuncher's 2007 research resolved time misalignment to under 10μs.


In birds


In invertebrates

Even though they do not have ears, invertebrates have developed other structures and systems to decode vibrations traveling through the air, or “sound”. Charles Henry Turner was the first scientist to formally show this phenomenon through rigorously controlled experiments in ants. Turner ruled out the detection of ground vibration and suggested that other insects likely have auditory systems as well. Many insects detect sound through the way air vibrations deflect hairs along their body. Some insects have even developed specialized hairs tuned to detecting particular frequencies, such as certain caterpillar species that have evolved hair with properties such that it resonates most with the sound of buzzing wasps, thus warning them of the presence of natural enemies.Tautz, Jürgen, and Michael Rostás. "Honeybee buzz attenuates plant damage by caterpillars." Current Biology 18, no. 24 (2008): R1125-R1126. Some insects possess a tympanal organ. These are "eardrums", that cover air filled chambers on the legs. Similar to the hearing process with vertebrates, the eardrums react to sonar waves. Receptors that are placed on the inside translate the oscillation into electric signals and send them to the brain. Several groups of flying insects that are preyed upon by echolocating bats can perceive the ultrasound emissions this way and reflexively practice ultrasound avoidance.


See also

; Basics * Ear * Hearing loss * Hearing test ; General * Audiology * Auditory scene analysis * Auditory science *
Auditory system The auditory system is the sensory system for the sense of hearing. It includes both the ear, sensory organs (the ears) and the auditory parts of the sensory system. System overview The outer ear funnels sound vibrations to the eardrum, incre ...
* Bone conduction * Hearing range * Human echolocation *
Listening Listening is the act of attention, paying attention to sounds. It includes listening to the sounds of Natural environment, nature, listening to music, and perhaps most importantly, Interpersonal communication, interpersonal listening, i.e. liste ...
* Neuronal encoding of sound *
Psychoacoustics Psychoacoustics is the branch of psychophysics involving the scientific study of the perception of sound by the human auditory system. It is the branch of science studying the psychological responses associated with sound including noise, speech, ...
*
Safe listening Safe listening is a Conceptual framework, framework for health promotion actions to ensure that sound-related recreational activities (such as Concert, concerts, nightclubs, and listening to music, broadcasts, or podcasts) do not pose a risk to ...
* Temporal envelope and fine structure * World Hearing Day ; Disorders *
Auditory processing disorder Auditory processing disorder (APD) is a neurodevelopmental disorder affecting the way the brain processes sounds. Individuals with APD usually have normal structure and function of the ear, but cannot process the information they hear in the s ...
* Endaural phenomena * Hearing loss * Hyperacusis * Presbycusis *
Tinnitus Tinnitus is a condition when a person hears a ringing sound or a different variety of sound when no corresponding external sound is present and other people cannot hear it. Nearly everyone experiences faint "normal tinnitus" in a completely ...
; Test and measurement * Audiogram * Audiometry * Auditory brainstem response (test) * Dichotic listening (test)


References


Further reading

*


External links

* Global Audiology- International Society of Audiology
World Health Organization, Deafness and Hearing Loss
* * *
Open University - OpenLearn - Article about hearing


NIOSH {{DEFAULTSORT:Hearing (Sense) Sound Audiology Auditory system Articles containing video clips