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System overview
The auditory system is the sensory system for the sense of hearing. It includes both the sensory organs
Auditory system
Structure (the ears) and the auditory parts of the sensory system.[1]
Anatomical terminology
Outer ear [edit on Wikidata]

Middle ear System overview [ edit ]

Inner ear The outer ear funnels sound vibrations to the eardrum, increasing the sound pressure in the middle
CC
Organ of Corti frequency range. The middle-ear ossicles further amplify the vibration pressure roughly 20 times.
The base of the stapes couples vibrations into the cochlea via the oval window, which vibrates the
Hair cell
perilymph liquid (present throughout the inner ear) and causes the round window to bulb out as the
Neurons oval window bulges in.[citation needed]
Neuronal structure
Vestibular and tympanic ducts are filled with perilymph, and the smaller cochlear duct between them 2:27
Cochlear nucleus is filled with endolymph, a fluid with a very different ion concentration and voltage.[2][3][4] Vestibular
How sounds make their way from the source to
Trapezoid body duct perilymph vibrations bend organ of Corti outer cells (4 lines) causing prestin to be released in
the brain
cell tips. This causes the cells to be chemically elongated and shrunk (somatic motor), and hair
Superior olivary complex
bundles to shift which, in turn, electrically affects the basilar membrane's movement (hair-bundle
Lateral lemniscus motor). These motors (outer hair cells) amplify the traveling wave amplitudes over 40-fold.[5] The outer hair cells (OHC) are minimally innervated by spiral
Inferior colliculi ganglion in slow (unmyelinated) reciprocal communicative bundles (30+ hairs per nerve fiber); this contrasts inner hair cells (IHC) that have only afferent
innervation (30+ nerve fibers per one hair) but are heavily connected. There are three to four times as many OHCs as IHCs. The basilar membrane (BM)
Medial geniculate nucleus
is a barrier between scalae, along the edge of which the IHCs and OHCs sit. Basilar membrane width and stiffness vary to control the frequencies best
Primary auditory cortex sensed by the IHC. At the cochlear base the BM is at its narrowest and most stiff (high-frequencies), while at the cochlear apex it is at its widest and least
The auditory ventral and stiff (low-frequencies). The tectorial membrane (TM) helps facilitate cochlear amplification by stimulating OHC (direct) and IHC (via endolymph
dorsal streams vibrations). TM width and stiffness parallels BM's and similarly aids in frequency differentiation.[6][7][8][9][10][11][12][13][14][excessive citations]
Clinical significance
The superior olivary complex (SOC), in the pons, is the first convergence of the left and right cochlear pulses. SOC has 14 described nuclei; their
See also abbreviation are used here (see Superior olivary complex for their full names). MSO determines the angle the sound came from by measuring time
References differences in left and right info. LSO normalizes sound levels between the ears; it uses the sound intensities to help determine sound angle. LSO
innervates the IHC. VNTB innervate OHC. MNTB inhibit LSO via glycine. LNTB are glycine-immune, used for fast signalling. DPO are high-frequency
and tonotopical. DLPO are low-frequency and tonotopical. VLPO have the same function as DPO, but act in a different area. PVO, CPO, RPO, VMPO,
ALPO and SPON (inhibited by glycine) are various signalling and inhibiting nuclei.[15][16][17][18]

The trapezoid body is where most of the cochlear nucleus (CN) fibers decussate (cross left to right and vice versa); this cross aids in sound
localization.[19] The CN breaks into ventral (VCN) and dorsal (DCN) regions. The VCN has three nuclei. [clarification needed] Bushy cells transmit timing info,
their shape averages timing jitters. Stellate (chopper) cells encode sound spectra (peaks and valleys) by spatial neural firing rates based on auditory
input strength (rather than frequency). Octopus cells have close to the best temporal precision while firing, they decode the auditory timing code. The
DCN has 2 nuclei. DCN also receives info from VCN. Fusiform cells integrate information to determine spectral cues to locations (for example, whether a
sound originated from in front or behind). Cochlear nerve fibers (30,000+) each have a most sensitive frequency and respond over a wide range of
levels.[20][21]

Simplified, nerve fibers' signals are transported by bushy cells to the binaural areas in the olivary complex, while signal peaks and valleys are noted by
stellate cells, and signal timing is extracted by octopus cells. The lateral lemniscus has three nuclei: dorsal nuclei respond best to bilateral input and have
complexity tuned responses; intermediate nuclei have broad tuning responses; and ventral nuclei have broad and moderately complex tuning curves.
Ventral nuclei of lateral lemniscus help the inferior colliculus (IC) decode amplitude modulated sounds by giving both phasic and tonic responses (short
and long notes, respectively). IC receives inputs not shown, including visual (pretectal area: moves eyes to sound. superior colliculus: orientation and
behavior toward objects, as well as eye movements (saccade)) areas, pons (superior cerebellar peduncle: thalamus to cerebellum connection/hear
sound and learn behavioral response), spinal cord (periaqueductal grey: hear sound and instinctually move), and thalamus. The above are what
implicate IC in the 'startle response' and ocular reflexes. Beyond multi-sensory integration IC responds to specific amplitude modulation frequencies,
allowing for the detection of pitch. IC also determines time differences in binaural hearing.[22] The medial geniculate nucleus divides into ventral (relay
and relay-inhibitory cells: frequency, intensity, and binaural info topographically relayed), dorsal (broad and complex tuned nuclei: connection to
somatosensory info), and medial (broad, complex, and narrow tuned nuclei: relay intensity and sound duration). The auditory cortex (AC) brings sound
into awareness/perception. AC identifies sounds (sound-name recognition) and also identifies the sound's origin location. AC is a topographical
frequency map with bundles reacting to different harmonies, timing and pitch. Right-hand-side AC is more sensitive to tonality, left-hand-side AC is more
sensitive to minute sequential differences in sound.[23][24] Rostromedial and ventrolateral prefrontal cortices are involved in activation during tonal space
and storing short-term memories, respectively.[25] The Heschl's gyrus/transverse temporal gyrus includes Wernicke's area and functionality, it is heavily
involved in emotion-sound, emotion-facial-expression, and sound-memory processes. The entorhinal cortex is the part of the 'hippocampus system' that
aids and stores visual and auditory memories.[26][27] The supramarginal gyrus (SMG) aids in language comprehension and is responsible for
compassionate responses. SMG links sounds to words with the angular gyrus and aids in word choice. SMG integrates tactile, visual, and auditory
info.[28][29]

Structure [ edit ]

Outer ear [ edit ]

Main article: Outer ear

The folds of cartilage surrounding the ear canal are called the pinna. Sound waves are reflected and attenuated
when they hit the pinna, and these changes provide additional information that will help the brain determine the
sound direction.

The sound waves enter the auditory canal, a deceptively simple tube. The ear canal amplifies sounds that are
between 3 and 12 kHz.[citation needed] The tympanic membrane, at the far end of the ear canal marks the
Anatomy of the human ear (The
beginning of the middle ear. length of the auditory canal is
exaggerated in this image.).
Middle ear [ edit ] Brown is outer ear.
Red is middle ear.
Main article: Middle ear Purple is inner ear.
Sound waves travel through the ear canal and hit the tympanic membrane, or eardrum. This wave information
travels across the air-filled middle ear cavity via a series of delicate bones: the malleus (hammer), incus (anvil)
and stapes (stirrup). These ossicles act as a lever, converting the lower-pressure eardrum sound vibrations into
higher-pressure sound vibrations at another, smaller membrane called the oval window or vestibular window. The
manubrium (handle) of the malleus articulates with the tympanic membrane, while the footplate (base) of the stapes
articulates with the oval window. Higher pressure is necessary at the oval window than at the tympanic membrane
because the inner ear beyond the oval window contains liquid rather than air. The stapedius reflex of the middle ear
muscles helps protect the inner ear from damage by reducing the transmission of sound energy when the stapedius
Auditory ossicles from a deep
muscle is activated in response to sound. The middle ear still contains the sound information in wave form; it is dissection of the tympanic cavity
converted to nerve impulses in the cochlea.

Inner ear [ edit ]

Main article: Inner ear

Cochlea
The inner ear consists of the cochlea and several non-auditory structures. The cochlea has three fluid-filled
sections (i.e. the scala media, scala tympani and scala vestibuli), and supports a fluid wave driven by
pressure across the basilar membrane separating two of the sections. Strikingly, one section, called the
cochlear duct or scala media, contains endolymph. The organ of Corti is located in this duct on the basilar
membrane, and transforms mechanical waves to electric signals in neurons. The other two sections are
known as the scala tympani and the scala vestibuli. These are located within the bony labyrinth, which is
filled with fluid called perilymph, similar in composition to cerebrospinal fluid. The chemical difference
Diagrammatic longitudinal section of the
between the fluids endolymph and perilymph fluids is important for the function of the inner ear due to cochlea. The cochlear duct, or scala media, is
electrical potential differences between potassium and calcium ions.[citation needed] labeled as ductus cochlearis at right.
Anatomical terminology
The plan view of the human cochlea (typical of all mammalian and most vertebrates) shows where specific
[edit on Wikidata]
frequencies occur along its length. The frequency is an approximately exponential function of the length of
the cochlea within the Organ of Corti. In some species, such as bats and dolphins, the relationship is expanded in specific areas to support their active
sonar capability.

Organ of Corti [ edit ]

Main article: Organ of Corti

The organ of Corti forms a ribbon of sensory epithelium which runs lengthwise down the cochlea's entire scala
media. Its hair cells transform the fluid waves into nerve signals. The journey of countless nerves begins with this
first step; from here, further processing leads to a panoply of auditory reactions and sensations.

Hair cell [ edit ]

Main article: Hair cell

Hair cells are columnar cells, each with a "hair bundle" of 100–200 specialized stereocilia at the top, for which they
are named. There are two types of hair cells specific to the auditory system; inner and outer hair cells. Inner hair
cells are the mechanoreceptors for hearing: they transduce the vibration of sound into electrical activity in nerve The organ of Corti located at the
scala media
fibers, which is transmitted to the brain. Outer hair cells are a motor structure. Sound energy causes changes in the
shape of these cells, which serves to amplify sound vibrations in a frequency specific manner. Lightly resting atop
the longest cilia of the inner hair cells is the tectorial membrane, which moves back and forth with each cycle of sound, tilting the cilia, which is what
elicits the hair cells' electrical responses.

Inner hair cells, like the photoreceptor cells of the eye, show a graded response, instead of the spikes typical of other neurons. These graded potentials
are not bound by the "all or none" properties of an action potential.

At this point, one may ask how such a wiggle of a hair bundle triggers a difference in membrane potential. The current model is that cilia are attached to
one another by "tip links", structures which link the tips of one cilium to another. Stretching and compressing, the tip links may open an ion channel and
produce the receptor potential in the hair cell. Recently it has been shown that cadherin-23 CDH23 and protocadherin-15 PCDH15 are the adhesion
molecules associated with these tip links.[30] It is thought that a calcium driven motor causes a shortening of these links to regenerate tensions. This
regeneration of tension allows for apprehension of prolonged auditory stimulation.[31]

Neurons [ edit ]
Main article: Hair cell neural connection

Afferent neurons innervate cochlear inner hair cells, at synapses where the neurotransmitter glutamate communicates signals from the hair cells to the
dendrites of the primary auditory neurons.

There are far fewer inner hair cells in the cochlea than afferent nerve fibers – many auditory nerve fibers innervate each hair cell. The neural dendrites
belong to neurons of the auditory nerve, which in turn joins the vestibular nerve to form the vestibulocochlear nerve, or cranial nerve number VIII.[32] The
region of the basilar membrane supplying the inputs to a particular afferent nerve fibre can be considered to be its receptive field.

Efferent projections from the brain to the cochlea also play a role in the perception of sound, although this is not well understood. Efferent synapses
occur on outer hair cells and on afferent (towards the brain) dendrites under inner hair cells

Neuronal structure [ edit ]

Cochlear nucleus [ edit ]

Main article: Cochlear nucleus

The cochlear nucleus is the first site of the neuronal processing of the newly converted "digital" data from the inner ear (see also binaural fusion). In
mammals, this region is anatomically and physiologically split into two regions, the dorsal cochlear nucleus (DCN), and ventral cochlear nucleus (VCN).
The VCN is further divided by the nerve root into the posteroventral cochlear nucleus (PVCN) and the anteroventral cochlear nucleus (AVCN).[33]

Trapezoid body [ edit ]

Main article: Trapezoid body

The trapezoid body is a bundle of decussating fibers in the ventral pons that carry information used for binaural computations in the brainstem. Some of
these axons come from the cochlear nucleus and cross over to the other side before traveling on to the superior olivary nucleus. This is believed to help
with localization of sound.[34]

Superior olivary complex [ edit ]

Main article: Superior olivary complex

The superior olivary complex is located in the pons, and receives projections predominantly from the ventral cochlear nucleus, although the dorsal
cochlear nucleus projects there as well, via the ventral acoustic stria. Within the superior olivary complex lies the lateral superior olive (LSO) and the
medial superior olive (MSO). The former is important in detecting interaural level differences while the latter is important in distinguishing interaural time
difference.[17]

Lateral lemniscus [ edit ]

Main article: Lateral lemniscus

The lateral lemniscus is a tract of axons in the brainstem that carries information about sound from the cochlear
nucleus to various brainstem nuclei and ultimately the contralateral inferior colliculus of the midbrain.

Inferior colliculi [ edit ]

Main article: Inferior colliculus

The inferior colliculi (IC) are located just below the visual processing centers known as the superior colliculi. The
central nucleus of the IC is a nearly obligatory relay in the ascending auditory system, and most likely acts to
integrate information (specifically regarding sound source localization from the superior olivary complex[16] and
dorsal cochlear nucleus) before sending it to the thalamus and cortex.[1] The inferior colliculus also receives
Lateral lemniscus in red, as it
descending inputs from the auditory cortex and auditory thalamus (or medial geniculate nucleus).[35]
connects the cochlear nucleus,
superior olivary nucleus and the
Medial geniculate nucleus [ edit ] inferior colliculus, seen from behind

Main article: Medial geniculate nucleus

The medial geniculate nucleus is part of the thalamic relay system.

Primary auditory cortex [ edit ]

Main article: Primary auditory cortex

The primary auditory cortex is the first region of cerebral cortex to receive auditory input.

Perception of sound is associated with the left posterior superior temporal gyrus (STG). The superior temporal gyrus contains several important
structures of the brain, including Brodmann areas 41 and 42, marking the location of the primary auditory cortex, the cortical region responsible for the
sensation of basic characteristics of sound such as pitch and rhythm. We know from research in nonhuman primates that the primary auditory cortex can
probably be divided further into functionally differentiable subregions.[36][37][38][39] [40][41][42] The neurons of the primary auditory cortex can be considered
to have receptive fields covering a range of auditory frequencies and have selective responses to harmonic pitches.[43] Neurons integrating information
from the two ears have receptive fields covering a particular region of auditory space.

The primary auditory cortex is surrounded by secondary auditory cortex, and interconnects with it. These secondary areas interconnect with further
processing areas in the superior temporal gyrus, in the dorsal bank of the superior temporal sulcus, and in the frontal lobe. In humans, connections of
these regions with the middle temporal gyrus are probably important for speech perception. The frontotemporal system underlying auditory perception
allows us to distinguish sounds as speech, music, or noise.

The auditory ventral and dorsal streams [ edit ]

Main article: Language processing in the brain

Further information: Two-streams hypothesis § Two auditory systems

From the primary auditory cortex emerge two separate pathways: the auditory ventral
stream and auditory dorsal stream.[44] The auditory ventral stream includes the anterior
superior temporal gyrus, anterior superior temporal sulcus, middle temporal gyrus and
temporal pole. Neurons in these areas are responsible for sound recognition, and extraction
of meaning from sentences. The auditory dorsal stream includes the posterior superior
temporal gyrus and sulcus, inferior parietal lobule and intra-parietal sulcus. Both pathways
project in humans to the inferior frontal gyrus. The most established role of the auditory
dorsal stream in primates is sound localization. In humans, the auditory dorsal stream in the
left hemisphere is also responsible for speech repetition and articulation, phonological long-
term encoding of word names, and verbal working memory.

Clinical significance [ edit ]

Proper function of the auditory system is required to able to sense, process, and
understand sound from the surroundings. Difficulty in sensing, processing and
understanding sound input has the potential to adversely impact an individual's ability to
communicate, learn and effectively complete routine tasks on a daily basis.[45]
Dual stream connectivity between the auditory cortex
and frontal lobe of monkeys and humans. Top: The auditory
In children, early diagnosis and treatment of impaired auditory system function is an cortex of the monkey (left) and human (right) is schematically
important factor in ensuring that key social, academic and speech/language developmental depicted on the supratemporal plane and observed from above
milestones are met.[46] (with the parieto- frontal operculi removed). Bottom: The brain
of the monkey (left) and human (right) is schematically
Impairment of the auditory system can include any of the following: depicted and displayed from the side. Orange frames mark the
region of the auditory cortex, which is displayed in the top sub-
Auditory brainstem response and ABR audiometry test for newborn hearing figures. Top and Bottom: Blue colors mark regions affiliated
Auditory processing disorder with the ADS, and red colors mark regions affiliated with the
AVS (dark red and blue regions mark the primary auditory
Hyperacusis fields). Abbreviations: AMYG-amygdala, HG-Heschl's gyrus,
Health effects due to noise FEF-frontal eye field, IFG-inferior frontal gyrus, INS-insula,
IPS-intra parietal sulcus, MTG-middle temporal gyrus, PC-pitch
Tinnitus
center, PMd-dorsal premotor cortex, PP-planum polare, PT-
Endaural phenomena planum temporale, TP-temporal pole, Spt-sylvian parietal-
temporal, pSTG/mSTG/aSTG-posterior/middle/anterior
See also [ edit ] superior temporal gyrus, CL/ ML/AL/RTL-
caudo-/middle-/antero-/rostrotemporal-lateral belt area,
Language processing in the brain CPB/RPB-caudal/rostral parabelt fields. Used with permission
from Poliva O. From where to what: a neuroanatomically
Neuroscience of music
based evolutionary model of the emergence of speech in
Selective auditory attention humans. Material was copied from this source, which
is available under a Creative Commons Attribution 4.0
International License .

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Further reading [ edit ]

Kandel, Eric R. (2012). Principles of Neural Science. New York: McGraw-Hill. ISBN 978-0-07-139011-8. OCLC 795553723 .

External links [ edit ]

Promenade 'round the cochlea Wikimedia Commons has

Auditory system – Washington University Neuroscience Tutorial media related to Auditory
system.
Lincoln Gray. "Chapter 13: Auditory System: Pathways and Reflexes" . Neuroscience Online, the Open-
Access Neuroscience Electronic Textbook. The University of Texas Health Science Center at Houston (UTHealth). Archived from the original on
2016-11-12. Retrieved 27 April 2014.

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