Wikipedia

Auditory system

Theauditory systemis thesensory systemfor the sense ofhearing.It includes both thesensory organs (the ears)and the auditory parts of thesensory system.[1]

Auditory system
Identifiers
FMA7192
Anatomical terminology
How sounds make their way from the source to the brain

System overview

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Theouter earfunnelssoundvibrations to theeardrum,increasing the sound pressure in the middle frequency range. Themiddle-earossiclesfurther amplify the vibration pressure roughly 20 times. The base of thestapescouples vibrations into thecochleavia theoval window,which vibrates theperilymphliquid (present throughout theinner ear) and causes theround windowto bulb out as the oval window bulges in.[1]

Vestibularandtympanic ductsare filled with perilymph, and the smallercochlear ductbetween them is filled withendolymph,a fluid with a very different ion concentration and voltage.[2][3][4]Vestibular duct perilymph vibrations bendorgan of Cortiouter cells (4 lines) causingprestinto be released in cell tips. This causes the cells to be chemically elongated and shrunk (somatic motor), and hair bundles to shift which, in turn, electrically affects thebasilar membrane's movement (hair-bundle motor). These motors (outerhair cells) amplify the traveling waveamplitudesover 40-fold.[5]The outer hair cells (OHC) are minimally innervated byspiral ganglionin slow (unmyelinated) reciprocal communicative bundles (30+ hairs pernerve fiber); this contrasts with 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.

Thebasilar membrane(BM) 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 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 stiff (low-frequencies). Thetectorial membrane(TM) helps facilitate cochlear amplification by stimulating OHC (direct) and IHC (via endolymph vibrations). TM width and stiffness parallels BM's and similarly aids in frequency differentiation.[6][7][8][9][10][11]

Thesuperior olivary complex(SOC), in thepons,is the first convergence of the left and right cochlear pulses. SOC has 14 described nuclei; their abbreviation are used here (seeSuperior olivary complexfor their full names). MSO determines the angle the sound came from by measuring time 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.[12][13][14][15]

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.[16]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.[17][18]

Simplified, nerve fibers' signals are transported by bushy cells to the binaural areas in theolivary 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:thalamustocerebellumconnection/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.[19]

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.[20][21]

Rostromedial and ventrolateral prefrontal cortices are involved in activation during tonal space and storing short-term memories, respectively.[22]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.[23][24]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.[25][26]

Structure

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Anatomy of the human ear (The length of the auditory canal is exaggerated in this image.).
Brown isouter ear.
Red ismiddle ear.
Purple isinner ear.

Outer ear

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Main article:Outer ear

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

The sound waves enter theauditory canal,a deceptively simple tube. The ear canal amplifies sounds that are between 3 and 12kHz.[1]Thetympanic membrane,at the far end of the ear canal marks the beginning of themiddle ear.

Middle ear

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Main article:Middle ear
Auditory ossicles from a deep dissection of the tympanic cavity

Sound waves travel through the ear canal and hit the tympanic membrane, oreardrum.This wave information travels across the air-filled middle ear cavity via a series of delicate bones: themalleus(hammer),incus(anvil) andstapes(stirrup). Theseossiclesact as a lever, converting the lower-pressure eardrum sound vibrations into higher-pressure sound vibrations at another, smaller membrane called theoval windowor vestibular window. Themanubrium(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. Thestapedius reflexof the middle ear muscles helps protect the inner ear from damage by reducing the transmission of sound energy when thestapedius muscleis activated in response to sound. The middle ear still contains the sound information in wave form; it is converted to nerve impulses in thecochlea.

Inner ear

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Cochlea
Diagrammatic longitudinal section of the cochlea. Thecochlear duct,orscala media,is labeled asductus cochlearisat right.
Identifiers
FMA7192
Anatomical terminology
Main article:Inner ear

The inner ear consists of thecochleaand several non-auditory structures. The cochlea has three fluid-filled sections (i.e. thescala media, scala tympani and scala vestibuli),and supports a fluid wave driven by pressure across thebasilar membraneseparating two of the sections. Strikingly, one section, called the cochlear duct orscala media,containsendolymph.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 thescala tympaniand thescala vestibuli.These are located within the bony labyrinth, which is filled with fluid calledperilymph,similar in composition to cerebrospinal fluid. The chemical difference between the fluidsendolymphandperilymphfluids is important for the function of the inner ear due to electrical potential differences between potassium and calcium ions.[citation needed]

The plan view of the human cochlea (typical of allmammalianand mostvertebrates) shows where specific frequencies occur along its length. The frequency is an approximately exponential function of the length of the cochlea within theOrgan 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

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Main article:Organ of Corti
Theorgan of Cortilocated at thescala media

The organ of Corti forms a ribbon of sensory epithelium which runs lengthwise down the cochlea's entirescala 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

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Main article:Hair cell

Hair cells are columnar cells, each with a "hair bundle" of 100–200 specializedstereociliaat the top, for which they are named. There are two types of hair cells specific to the auditory system;innerandouterhaircells.Inner hair cells are the mechanoreceptors for hearing: they transduce the vibration of sound into electrical activity innerve 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 longestciliaof the inner hair cells is thetectorial 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 thephotoreceptor cellsof the eye, show agraded response,instead of thespikestypical 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 thatcadherin-23 CDH23andprotocadherin-15 PCDH15are the adhesion molecules associated with these tip links.[27]It is thought that acalciumdriven motor causes a shortening of these links to regenerate tensions. This regeneration of tension allows for apprehension of prolonged auditory stimulation.[28]

Neurons

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Main article:Hair cell neural connection

Afferent neurons innervate cochlear inner hair cells, at synapses where the neurotransmitterglutamatecommunicates 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 theauditory nerve,which in turn joins thevestibular nerveto form thevestibulocochlear nerve,orcranial nervenumber VIII.[29] The region of the basilar membrane supplying the inputs to a particular afferent nerve fibre can be considered to be itsreceptive 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

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Cochlear nucleus

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Main article:Cochlear nucleus

Thecochlear nucleusis the first site of the neuronal processing of the newly converted "digital" data from the inner ear (see alsobinaural fusion). In mammals, this region is anatomically and physiologically split into two regions, thedorsal cochlear nucleus(DCN), andventral cochlear nucleus(VCN). The VCN is further divided by the nerve root into the posteroventral cochlear nucleus (PVCN) and the anteroventral cochlear nucleus (AVCN).[30]

Trapezoid body

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Main article:Trapezoid body

Thetrapezoid bodyis a bundle ofdecussatingfibers in the ventral pons that carry information used for binaural computations in the brainstem. Some of theseaxonscome from thecochlear nucleusandcross overto the other side before traveling on to thesuperior olivary nucleus.This is believed to help withlocalization of sound.[31]

Superior olivary complex

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Main article:Superior olivary complex

Thesuperior olivary complexis located in thepons,and receives projections predominantly from the ventral cochlear nucleus, although the dorsal cochlear nucleus projects there as well, via the ventral acoustic stria. Within thesuperior olivary complexlies 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 distinguishinginteraural time difference.[14]

Lateral lemniscusin red, as it connects thecochlear nucleus,superior olivary nucleusand theinferior colliculus,seen from behind

Lateral lemniscus

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Main article:Lateral lemniscus

The lateral lemniscus is a tract ofaxonsin thebrainstemthat carries information about sound from thecochlear nucleusto various brainstem nuclei and ultimately the contralateralinferior colliculusof themidbrain.

Inferior colliculi

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Main article:Inferior colliculus

The inferior colliculi (IC) are located just below the visual processing centers known as thesuperior 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 thesuperior olivary complex[13]anddorsal cochlear nucleus) before sending it to thethalamusandcortex.[1]The inferior colliculus also receives descending inputs from theauditory cortexand auditorythalamus(ormedial geniculate nucleus).[32]

Medial geniculate nucleus

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Main article:Medial geniculate nucleus

Themedial geniculate nucleusis part of the thalamic relay system.

Primary auditory cortex

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Main article:Primary auditory cortex

Theprimary auditory cortexis the first region ofcerebral cortexto receive auditory input.

Perception of sound is associated with the left posteriorsuperior temporal gyrus(STG). The superior temporal gyrus contains several important structures of the brain, includingBrodmann areas41 and 42, marking the location of theprimary 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.[33][34][35][36] [37][38][39] The neurons of the primary auditory cortex can be considered to havereceptive fieldscovering a range ofauditory frequenciesand have selective responses to harmonic pitches.[40]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 thesuperior temporal gyrus,in the dorsal bank of thesuperior temporal sulcus,and in thefrontal lobe.In humans, connections of these regions with themiddle temporal gyrusare probably important forspeech perception.The frontotemporal system underlying auditory perception allows us to distinguish sounds as speech, music, or noise.

The auditory ventral and dorsal streams

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Main article:Language processing in the brain
Further information:Two-streams hypothesis § Two auditory systems
Dual stream connectivity between the auditory cortex and frontal lobe of monkeys and humans.Top: The auditory cortex of the monkey (left) and human (right) is schematically depicted on the supratemporal plane and observed from above (with the parieto- frontal operculi removed). Bottom: The brain of the monkey (left) and human (right) is schematically depicted and displayed from the side. Orange frames mark the region of the auditory cortex, which is displayed in the top sub-figures. Top and Bottom: Blue colors mark regions affiliated with the ADS, and red colors mark regions affiliated with the AVS (dark red and blue regions mark the primary auditory fields). Abbreviations: AMYG-amygdala, HG-Heschl's gyrus, FEF-frontal eye field, IFG-inferior frontal gyrus, INS-insula, IPS-intra parietal sulcus, MTG-middle temporal gyrus, PC-pitch center, PMd-dorsal premotor cortex, PP-planum polare, PT-planum temporale, TP-temporal pole, Spt-sylvian parietal-temporal, pSTG/mSTG/aSTG-posterior/middle/anterior superior temporal gyrus, CL/ ML/AL/RTL-caudo-/middle-/antero-/rostrotemporal-lateral belt area, CPB/RPB-caudal/rostral parabelt fields. Used with permission from Poliva O. From where to what: a neuroanatomically based evolutionary model of the emergence of speech in humans.Material was copied from this source, which is available under aCreative Commons Attribution 4.0 International License.

From the primary auditory cortex emerge two separate pathways: the auditory ventral stream and auditory dorsal stream.[41]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 lobuleand 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

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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.[42]

In children, early diagnosis and treatment of impaired auditory system function is an important factor in ensuring that key social, academic and speech/language developmental milestones are met.[43]

Impairment of the auditory system can include any of the following:

See also

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References

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