Stereocilia (inner ear)

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This article is about the stereocilia of the ear. For the stereocilia of the epididymis, see Stereocilia (epididymis).
See also: Hair cell
Section through the spiral organ of Corti, magnified. The stereocilia are the "hairs" sticking out of the tops of the inner and outer hair cells.
Stereocilia of frog inner ear.

In the inner ear, stereocilia are the mechanosensing organelles of hair cells, which respond to fluid motion in numerous types of animals for various functions, including hearing and balance. They are about 10–50 micrometers in length and share some similar features of microvilli.[1] The hair cells turn the fluid pressure and other mechanical stimuli into electric stimuli via the many microvilli that make up stereocilia rods.[2] Stereocilia exist in the auditory and vestibular systems.

In the Auditory Pathway[edit]

As acoustic sensors in mammals, stereocilia are lined up in the Organ of Corti within the cochlea of the inner ear. In hearing, stereocilia transform the mechanical energy of sound waves into electrical signals for the hair cells, which ultimately leads to an excitation of the auditory nerve. Stereocilia are composed of cytoplasm with embedded bundles of cross-linked actin filaments. The actin filaments anchor to the terminal web and the top of the cell membrane and are arranged in grade of height.[2] When the stapes causes sound waves in the endolymphatic fluid in the cochlea, the stereocilia are deflected in a shearing motion, which results in the mentioned electrical signal for the hair cell.

In the Vestibular Pathway[edit]

In the vestibular system, the stereocilia are located in the otolithic organs and the semicircular canals. Hair cells in the vestibular system are slightly different from those in the auditory system, in that vestibular hair cells have one tallest cilium, termed the kinocilium. Bending the stereocilia toward the kinocilium depolarizes the cell and results in increased afferent activity. Bending the stereocilia away from the kinocilium hyperpolarizes the cell and results in a decrease in afferent activity. In the semicircular canals, the hair cells are found in the crista ampullaris, and the stereocilia protrude into the ampullary cupula. Here, the stereocilia are all oriented in the same direction. In the otoliths, the hair cells are topped by small, calcium carbonate crystals called otoconia. Unlike the semicircular ducts, the kinocilia of hair cells in the otoliths are not oriented in a consistent direction. The kinocilia point toward (in the utricle) or away from (in the saccule) a middle line called the striola.[3]

Design and constellation[edit]

Stereociliar design and constellation is important for mechanoelectrical transduction. Resembling hair-like projections, the stereocilia are arranged in bundles of 30-300.[4] Within the bundles the stereocilia are often lined up in several rows of increasing height, similar to a staircase. At the core of these hair-like stereocilia are rigid cross-linked actin filaments, which can renew every 48 hours. These actin filaments face their positive ends at the tips of the stereocilia and their negative ends at the base and can be up to 120 micrometres in length.[4] Filamentous structures, called tip links, connect the tips of stereocilia in adjacent rows in the bundles. The tip links are made up of nearly vertical fine filaments that run upward from the top end of a shorter stereocilia to its taller neighbor.[2] Tip links are analogous to tiny springs, which, when stretched, open cation selective channels thus allowing ions to flow across the cell membrane into the hair cells. They also are involved in the force transmission across the bundle and the maintenance of the hair bundle structure.[5]

Mechanoelectrical transduction[edit]

In the cochlea, a shearing movement between the tectorial membrane and the basilar membrane deflects the stereocilia, affecting the tension on the tip-link filaments, which then open and close the non-specific ion channels.[2] When tension increases, the flow of ions across the membrane into the hair cell rises as well. Such influx of ions causes a depolarization of the cell, resulting in an electrical potential that ultimately leads to a signal for the auditory nerve and the brain. The identity of the mechanosensitive channels in the stereocilia is still unknown.

The transduction channels associated with stereocilia are thought to lie at the distal ends of the stereocilia.[6] Deflections of the stereocilia in the direction of the tallest stereocilia leads to an increased rate of opening of nonspecific cation channels. This, in turn, causes receptor depolarization and leads to the excitement of the cochlear nerve afferents that are located at the base of the hair cell. Deflections of the stereocilia in the opposite direction toward the shortest stereocilia causes transduction channels to close. In this situation, the hair cells become hyperpolarized and the nerve afferents are not excited.[7][8][9]

There are two different types of fluid that surround the hair cells of the inner ear. The endolymph is the fluid that surrounds the apical surfaces of hair cells. Potassium is the major cation in the endolymph and is thought to be responsible for carrying the receptor currents in the cochlea. Perilymph is found surrounding the sides and the bases of the hair cells. Perilymph is low in potassium and high in sodium.[10][11] The different ionic makeups of the surrounding fluid in addition to the resting potential of the hair cell creates a potential difference across the apical membrane of the hair cell, so potassium enters when transduction channels open. An influx of potassium ions depolarizes the cell and causes the release of a neurotransmitter that can initiate nerve impulses in the sensory neurons that synapse on the base of the hair cell.

Destruction of stereocilia[edit]

Stereocilia (along with the entirety of the hair cell) in mammals can be damaged or destroyed by excessive loud noises, disease, and toxins and are not regenerable.[4] Environmental noise induced hearing impairment is probably the most prevalent noise health effect according to the U.S. Environmental Protection Agency. Abnormal structure/organization of a bundle of stereocilia can also cause deafness and in turn create balance problems for an individual. In other vertebrates, if the hair cell is harmed, supporting cells will divide and replace the damaged hair cells.[2]

References[edit]

  1. ^ Caceci, T. VM8054 Veterinary Histology: Male Reproductive System. http://education.vetmed.vt.edu/Curriculum/VM8054/Labs/Lab27/Lab27.htm (accessed 2/16/06).
  2. ^ a b c d e Alberts, B., Johnson, A., Lewis, J., Raff, M., Roberts, K. and Walter, P. (2002) The Molecular Biology of the Cell. Garland Science Textbooks.
  3. ^ Gray, Lincoln. "Vestibular System: Structure and Function". Neuroscience Online: an electronic book for the neurosciences. http://education.vetmed.vt.edu/Curriculum/VM8054/Labs/Lab27/Lab27.htm (accessed 2/16/06).
  4. ^ a b c Rzadzinska AK, Schneider ME, Davies C, Riordan GP, Kachar B (2004). "An actin molecular treadmill and myosins maintain stereocilia functional architecture and self-renewal". J. Cell Biol. 164 (6): 887–97. doi:10.1083/jcb.200310055. PMC 2172292. PMID 15024034. 
  5. ^ Tsuprun V, Santi P (2002). "Structure of outer hair cell stereocilia side and attachment links in the chinchilla cochlea.". J. Histochem. Cytochem. 50 (4): 493–502. doi:10.1177/002215540205000406. PMID 11897802. 
  6. ^ Hudspeth, A. J. (1982). "Extracellular current flow and the site of transduction by vertebrate hair cells". The Journal of neuroscience : the official journal of the Society for Neuroscience 2 (1): 1–10. PMID 6275046.  edit
  7. ^ Hackney, C. M.; Furness, D. N. (1995). "Mechanotransduction in vertebrate hair cells: Structure and function of the stereociliary bundle". The American journal of physiology 268 (1 Pt 1): C1–13. PMID 7840137.  edit
  8. ^ Corey, D. P.; Hudspeth, A. J. (1979). "Ionic basis of the receptor potential in a vertebrate hair cell". Nature 281 (5733): 675–677. doi:10.1038/281675a0. PMID 45121.  edit
  9. ^ Ohmori, H. (1985). "Mechano-electrical transduction currents in isolated vestibular hair cells of the chick". The Journal of physiology 359: 189–217. PMC 1193371. PMID 2582113.  edit
  10. ^ Corey, D. P.; Hudspeth, A. J. (1979). "Ionic basis of the receptor potential in a vertebrate hair cell". Nature 281 (5733): 675–677. doi:10.1038/281675a0. PMID 45121.  edit
  11. ^ Bosher, S. K.; Warren, R. L. (1978). "Very low calcium content of cochlear endolymph, an extracellular fluid". Nature 273 (5661): 377–378. doi:10.1038/273377a0. PMID 661948.  edit