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Hearing, or auditory perception, is the ability to perceive sounds through an organ, such as an ear, by detecting vibrations 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,
liquid A liquid is a nearly incompressible fluid that conforms to the shape of its container but retains a (nearly) constant volume independent of pressure. As such, it is one of the four fundamental states of matter (the others being solid, gas, a ...
, or gaseous matter. It is one of the traditional five senses. 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 sensory organs (the ears) and the auditory parts of the sensory system. System overview The outer ear funnels sound vibrations to the eardrum, increasin ...
: mechanical waves, known as vibrations, are detected by the ear and transduced into nerve impulses that are perceived by the brain (primarily in the temporal lobe). Like touch, 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 sensory organs (the ears) and the auditory parts of the sensory system. System overview The outer ear funnels sound vibrations to the eardrum, increasin ...
: 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 pinna to the eardrum and is about in length and in diameter. Struc ...
, which terminates at the
eardrum In the anatomy of humans and various other tetrapods, the eardrum, also called the tympanic membrane or myringa, is a thin, cone-shaped membrane that separates the external ear The outer ear, external ear, or auris externa is the extern ...
, 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 of the sound. Cerumen (ear wax) is produced by
ceruminous Ceruminous glands are specialized sudoriferous glands (sweat glands) located subcutaneously in the external auditory canal, in the outer 1/3. Ceruminous glands are simple, coiled, tubular glands made up of an inner secretory layer of cells and an ou ...
and sebaceous glands 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 The ossicles (also called auditory ossicles) are three bones in either middle ear that are among the smallest bones in the human body. They serve to transmit sounds from the air to the fluid-filled labyrinth (cochlea). The absence of the auditory ...
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 purpose of the middle ear ossicles is to overcome the impedance mismatch between air waves and cochlear waves, by providing impedance matching. 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, 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 In physiology, tonotopy (from Greek tono = frequency and topos = place) is the spatial arrangement of where sounds of different frequency are processed in the brain. Tones close to each other in terms of frequency are represented in topologically ...
, 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 of the hair cells, specialized auditory receptors located within the organ of Corti. While the hair cells do not produce action potentials themselves, they release neurotransmitter at synapses with the fibers of the
auditory nerve The cochlear nerve (also auditory nerve or acoustic nerve) is one of two parts of the vestibulocochlear nerve, a cranial nerve present in amniotes, the other part being the vestibular nerve. The cochlear nerve carries auditory sensory information ...
, which does produce action potentials. In this way, the patterns of oscillations on the basilar membrane are converted to
spatiotemporal pattern Spatiotemporal patterns are patterns that occur in a wide range of natural phenoma and are characterized by a spatial and a temporal patterning. The general rules of pattern formation hold. In contrast to "static", pure spatial patterns, the ...
s 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 cont ...
.


Neuronal

The sound information from the cochlea travels via the
auditory nerve The cochlear nerve (also auditory nerve or acoustic nerve) is one of two parts of the vestibulocochlear nerve, a cranial nerve present in amniotes, the other part being the vestibular nerve. The cochlear nerve carries auditory sensory information ...
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 cont ...
. From there, the signals are projected to the inferior colliculus in the midbrain tectum. The inferior colliculus 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 stimuli, such as sudden noise or sharp movement, and is associated with negative Affect (psychology), affect.Rammirez-Moreno, D ...
. The inferior colliculus in turn projects to the medial geniculate nucleus, a part of the thalamus where sound information is relayed to the
primary auditory cortex The auditory cortex is the part of the temporal lobe that processes auditory information in humans and many other vertebrates. It is a part of the auditory system, performing basic and higher functions in hearing, such as possible relations to ...
in the temporal lobe. Sound is believed to first become consciously experienced at the
primary auditory cortex The auditory cortex is the part of the temporal lobe that processes auditory information in humans and many other vertebrates. It is a part of the auditory system, performing basic and higher functions in hearing, such as possible relations to ...
. Around the
primary auditory cortex The auditory cortex is the part of the temporal lobe that processes auditory information in humans and many other vertebrates. It is a part of the auditory system, performing basic and higher functions in hearing, such as possible relations to ...
lies
Wernickes area Wernicke's area (; ), also called Wernicke's speech area, is one of the two parts of the cerebral cortex that are linked to speech, the other being Broca's area. It is involved in the comprehension of written and spoken language, in contrast to B ...
, a cortical area involved in interpreting sounds that is necessary to understand spoken words. Disturbances (such as
stroke A stroke is a medical condition in which poor blood flow to the brain causes cell death. There are two main types of stroke: ischemic, due to lack of blood flow, and hemorrhagic, due to bleeding. Both cause parts of the brain to stop functionin ...
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 An auditory hallucination, or paracusia, is a form of hallucination that involves perceiving sounds without auditory stimulus. While experiencing an auditory hallucination, the affected person would hear a sound or sounds which did not come from t ...
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 Hearing, or auditory perception, is the ability to perceive sounds through an organ, such as an ear, by detecting vibrations as periodic changes in the pressure of a surrounding medium. The academic field concerned with hearing is auditory ...
. 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 or sensory organ (cochlea and associated structures) or the vestibulocochlear nerve (cranial nerve VIII). SNHL accounts for about 90% of rep ...
and mixed types. 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 Post-lingual deafness is a deafness which develops after the acquisition of speech and language, usually after the age of six. Post-lingual hearing impairments are far more common than prelingual deafness. Typically, hearing loss is gradual, and of ...
. 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, which absorbs nearly all sound. Another means is the use of devices such as earplugs, which are inserted into the ear canal to block noise, or
earmuffs Earmuffs are clothing accessories or personal protective equipment designed to cover a person's ears for hearing protection or warmth. They consist of a thermoplastic or metal head-band that fits over the top or back of the head, and a cushion o ...
, 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 neurodegeneration, neurodegenerative disease that usually starts slowly and progressively worsens. It is the cause of 60–70% of cases of dementia. The most common early symptom is difficulty in short-term me ...
and dementia 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. 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 Audio most commonly refers to sound, as it is transmitted in signal form. It may also refer to: Sound *Audio signal, an electrical representation of sound *Audio frequency, a frequency in the audio spectrum *Digital audio, representation of sound ...
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 mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". 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 bi ...
use ultrasound for echolocation while in flight. Dogs are able to hear ultrasound, which is the principle of 'silent'
dog whistle A dog whistle (also known as silent whistle or Galton's whistle) is a type of whistle that emits sound in the ultrasonic range, which humans cannot hear but some other animals can, including dogs and domestic cats, and is used in their training ...
s. Snakes sense infrasound through their jaws, and baleen whales,
giraffe The giraffe is a large African hoofed mammal belonging to the genus ''Giraffa''. It is the tallest living terrestrial animal and the largest ruminant on Earth. Traditionally, giraffes were thought to be one species, ''Giraffa camelopardalis ...
s, dolphins and elephants use it for communication. Some fish 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 Carp are various species of oily freshwater fish from the family Cyprinidae, a very large group of fish native to Europe and Asia. While carp is consumed in many parts of the world, they are generally considered an invasive species in parts of ...
and
herring Herring are forage fish, mostly belonging to the family of Clupeidae. Herring often move in large schools around fishing banks and near the coast, found particularly in shallow, temperate waters of the North Pacific and North Atlantic Oceans, i ...
.


In invertebrates

Even though they don’t 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 Ultrasound avoidance is an escape or avoidance reflex displayed by certain animal species that are preyed upon by echolocating predators. Ultrasound avoidance is known for several groups of insects that have independently evolved mechanisms for ult ...
.


See also


Physiological

* Ear * Hearing loss * Hearing test


General

*
Auditory scene analysis In perception and psychophysics, auditory scene analysis (ASA) is a proposed model for the basis of auditory perception. This is understood as the process by which the human auditory system organizes sound into perceptually meaningful elements. T ...
* Auditory science *
Auditory system The auditory system is the sensory system for the sense of hearing. It includes both the sensory organs (the ears) and the auditory parts of the sensory system. System overview The outer ear funnels sound vibrations to the eardrum, increasin ...
* Bone conduction * Hearing range * Human echolocation * Listening *
Neuronal encoding of sound The neural encoding of sound is the representation of auditory sensation and perception in the nervous system. This article explores the basic physiological principles of sound perception, and traces hearing mechanisms from sound as pressure wa ...
* Temporal envelope and fine structure *
World Hearing Day World Hearing Day is a campaign held each year by Office of Prevention of Blindness and Deafness of the World Health Organization (WHO). Activities take place across the globe and an event is hosted at the World Health Organization on March 3. The ...
*
Safe listening Safe listening is a framework for health promotion actions to ensure that sound-related recreational activities (such as concerts, nightclubs, and listening to music, broadcasts, or podcasts) do not pose a risk to hearing. While research shows ...


Test and measurement

* Audiogram * Audiometry *
Dichotic listening Dichotic listening is a psychological test commonly used to investigate selective attention and the lateralization of brain function within the auditory system. It is used within the fields of cognitive psychology and neuroscience. In a standar ...
(test) * Auditory brainstem response (test)


Disorders

* Auditory processing disorder *
Endaural phenomena Endaural phenomena are sounds that are heard without any external acoustic stimulation. Endaural means "in the ear". Phenomena include transient ringing in the ears (that sound like sine tones), white noise-like sounds, and subjective tinnitus. En ...
* Hearing loss * Hyperacusis * Presbycusis * Tinnitus


References


Further reading

*


External links


World Health Organization, Deafness and Hearing Loss
* * *
Open University - OpenLearn - Article about hearing
{{DEFAULTSORT:Hearing (Sense) Sound Audiology Auditory system