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Neuron: Hair cell
Section through the spiral organ of Corti. Magnified. ("Outer hair cells" labeled near top; "inner hair cells" labeled near center).
Location	Cochlea
Function	Amplify sound waves and transduce auditory information to the Brain Stem
Morphology	Unique (see text)
Presynaptic connections	None
Postsynaptic connections	Via auditory nerve to vestibulocochlear nerve to inferior colliculus
Gray's	subject #232 1057
NeuroLex ID	nifext_61
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Hair cells are the sensory receptors of both the auditory system and the vestibular system in all vertebrates. In mammals, the auditory hair cells are located within the organ of Corti on a thin basilar membrane in the cochlea of the inner ear. They derive their name from the tufts of stereocilia that protrude from the apical surface of the cell, a structure known as the hair bundle, into the scala media, a fluid-filled tube within the cochlea. Mammalian cochlear hair cells come in two anatomically and functionally distinct types: the outer and inner hair cells. Damage to these hair cells results in decreased hearing sensitivity, i.e. sensorineural hearing loss.

[edit] Hair bundles as sound detectors and amplifiers

Research of the past decades has shown that outer hair cells do not send neural signals to the brain, but that they mechanically amplify low-level sound that enters the cochlea. The amplification may be powered by movement of their hair bundles, or by an electrically driven motility of their cell bodies. The inner hair cells transform the sound vibrations in the fluids of the cochlea into electrical signals that are then relayed via the auditory nerve to the auditory brainstem and to the auditory cortex.

Results in recent years further indicate that mammals apparently have conserved an evolutionarily earlier type of hair-cell motility. This so-called hair-bundle motility amplifies sound in all non-mammalian land vertebrates. It is affected by the closing mechanism of the mechanical sensory ion channels at the tips of the hair bundles. Thus, the same hair-bundle mechanism that detects sound vibrations also actively “vibrates back” and thereby mechanically amplifies weak incoming sound.

[edit] Inner hair cells – from sound to nerve signal

Section through the organ of corti, showing position of inner and outer hair cells

The deflection of the hair-cell stereocilia opens mechanically gated ion channels that allow any small, positively charged ions (primarily potassium and calcium) to enter the cell. Unlike many other electrically active cells, the hair cell itself does not fire an action potential. Instead, the influx of positive ions from the endolymph in Scala media depolarizes the cell, resulting in a receptor potential. This receptor potential opens voltage gated calcium channels; calcium ions then enter the cell and trigger the release of neurotransmitters at the basal end of the cell. The neurotransmitters diffuse across the narrow space between the hair cell and a nerve terminal, where they then bind to receptors and thus trigger action potentials in the nerve. In this way, the mechanical sound signal is converted into an electrical nerve signal. The repolarization in the hair cell is done in a special manner. The perilymph in Scala tympani has a very low concentration of positive ions. The electrochemical gradient makes the positive ions flow through channels to the perilymph.

Hair cells chronically leak Ca⁺². This leakage causes a tonic release of neurotransmitter to the synapses. It is thought that this tonic release is what allows the hair cells to respond so quickly in response to mechanical stimuli. The quickness of the hair cell response may also be due to that fact that it can increase the amount of neurotransmitter release in response to a change as little as 100 μV in membrane potential.^[1]

[edit] Outer hair cells – acoustical pre-amplifiers

In mammalian outer hair cells, the receptor potential triggers active vibrations of the cell body. This mechanical response to electrical signals is termed somatic electromotility^[2] and drives oscillations in the cell’s length, which occur at the frequency of the incoming sound and in a stable phase relation. Outer hair cells have evolved only in mammals. They have not improved hearing sensitivity, which reaches similarly exquisite values also in other classes of vertebrates. But they have extended the hearing range from about 11 kHz in some types of bird to about 200 kHz in some marine mammals. They have also improved frequency selectivity (frequency discrimination), which is of particular benefit for humans, because it enabled sophisticated speech and music.

The molecular biology of hair cells has seen considerable progress in recent years, with the identification of the motor protein (prestin) that underlies somatic electromotility in the outer hair cells. Santos-Sacchi et al. have shown that prestin's function is dependent on chloride channel signalling and that it is compromised by the common marine pesticide tributyltin (TBT). Because this class of pollutant bioconcentrates up the food chain, the effect is pronounced in top marine predators such as Orcas and toothed whales.^[3]

[edit] Neural connection

Neurons of the auditory or vestibulocochlear nerve (the VIIIth cranial nerve) innervate cochlear and vestibular hair cells.^[4] The neurotransmitter released by hair cells to stimulate the dendrites of afferent neurons is thought to be glutamate. At the presynaptic juncture, there is a distinct presynaptic dense body or ribbon. This dense body is surrounded by synaptic vesicles and is thought to aid in the fast release of neurotransmitter.

Nerve fiber innervation is much denser for inner hair cells than for outer hair cells. A single inner hair cell is innervated by numerous nerve fibers, whereas a single nerve fiber innervates many outer hair cells. Inner hair cell nerve fibers are also very heavily myelinated, which is in contrast to the unmyelinated outer hair cell nerve fibers.

Efferent projections from the brain to the cochlea also play a role in the perception of sound. Efferent synapses occur on outer hair cells and on afferent (towards the brain) dendrites under inner hair cells. The presynaptic terminal bouton is filled with vesicles containing acetylcholine and a neuropeptide called Calcitonin gene-related peptide (CGRP). The effects of these compounds varies, in some hair cells the acetylcholine hyperpolarized the cell, which reduces the sensitivity of the cochlea locally.

[edit] Additional images

The lamina reticularis and subjacent structures.

Inner ear illustration showing semicircular canal, hair cells, ampulla, cupula, vestibular nerve, & fluid

[edit] References

1. ^ Chan DK, Hudspeth AJ (2005 Feb). "Ca2+ current-driven nonlinear amplification by the mammalian cochlea in vitro". Nature Neuroscience 8 (2): 149–155. doi:10.1038/nn1385. PMID 15643426. 
2. ^ Brownell WE, Bader CR, Bertrand D, de Ribaupierre Y (1985-01-11). "Evoked mechanical responses of isolated cochlear outer hair cells". Science 227 (4683): 194-196. doi:10.1126/science.3966153. PMID 3966153. 
3. ^ Santos-Sacchi Joseph, Song Lei, Zheng Jiefu, Nuttall Alfred L (2006-04-12). "Control of Mammalian Cochlear Amplification by Chloride Anions". Journal of Neuroscience 26 (15): 3992–3998. doi:10.1523/JNEUROSCI.4548-05.2006. PMID 16611815. http://www.jneurosci.org/cgi/content/full/26/15/3992. 
4. ^ "Cranial Nerve VIII. Vestibulocochlear Nerve". Meddean. http://www.meddean.luc.edu/lumen/meded/GrossAnatomy/h_n/cn/cn1/cn8.htm. Retrieved 2008-06-04. 

• Manley GA, Popper AN, Fay RR (2004). Evolution of the Vertebrate Auditory System. New York: Springer-Verlag. ISBN 0387210938. 
• Kandel ER, Schwartz JH, Jessell TM (2000). Principles of Neural Science (4th ed.). New York: McGraw-Hill. pp. 590–594. ISBN 0838577016. 
• Coffin A, Kelley M, Manley GA, Popper AN. "Evolution of sensory hair cells". pp. 55–94.  in Manley et al. (2004)
• Manley GA. "Advances and perspectives in the study of the evolution of the vertebrate auditory system". pp. 360–368.  in Manley et al. (2004)
• Fettiplace R, Hackney CM (2006). "The sensory and motor roles of auditory hair cells". Nature Reviews. Neuroscience 7 (1): 19–29. doi:10.1038/nrn1828. PMID 16371947. 

[edit] External links

• Hair Cells at the University of Montpellier
• Dancing OHC video Yale Ear Lab
• NIF Search - Hair Cell via the Neuroscience Information Framework