Auditory Ossicles
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The ossicles (also called auditory ossicles) are three
bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, ...
s in either
middle ear The middle ear is the portion of the ear medial to the eardrum, and distal to the oval window of the cochlea (of the inner ear). The mammalian middle ear contains three ossicles, which transfer the vibrations of the eardrum into waves in the ...
that are among the smallest bones in the human body. They serve to transmit
sound In physics, sound is a vibration that propagates as an acoustic wave, through a transmission medium such as a gas, liquid or solid. In human physiology and psychology, sound is the ''reception'' of such waves and their ''perception'' b ...
s from the air to the fluid-filled
labyrinth In Greek mythology, the Labyrinth (, ) was an elaborate, confusing structure designed and built by the legendary artificer Daedalus for King Minos of Crete at Knossos. Its function was to hold the Minotaur, the monster eventually killed by t ...
(
cochlea The cochlea is the part of the inner ear involved in hearing. It is a spiral-shaped cavity in the bony labyrinth, in humans making 2.75 turns around its axis, the modiolus. A core component of the cochlea is the Organ of Corti, the sensory or ...
). The absence of the auditory ossicles would constitute a moderate-to-severe
hearing loss Hearing loss is a partial or total inability to hear. Hearing loss may be present at birth or acquired at any time afterwards. Hearing loss may occur in one or both ears. In children, hearing problems can affect the ability to acquire spoken ...
. The term "ossicle" literally means "tiny bone". Though the term may refer to any small bone throughout the body, it typically refers to the
malleus The malleus, or hammer, is a hammer-shaped small bone or ossicle of the middle ear. It connects with the incus, and is attached to the inner surface of the eardrum. The word is Latin for 'hammer' or 'mallet'. It transmits the sound vibrations f ...
,
incus The ''incus'' (plural incudes) or anvil is a bone in the middle ear. The anvil-shaped small bone is one of three ossicles in the middle ear. The ''incus'' receives vibrations from the ''malleus'', to which it is connected laterally, and transmit ...
, and
stapes The ''stapes'' or stirrup is a bone in the middle ear of humans and other animals which is involved in the conduction of sound vibrations to the inner ear. This bone is connected to the oval window by its annular ligament, which allows the foo ...
(hammer, anvil, and stirrup) of the middle
ear An ear is the organ that enables hearing and, in mammals, body balance using the vestibular system. In mammals, the ear is usually described as having three parts—the outer ear, the middle ear and the inner ear. The outer ear consists of ...
.


Structure

The ossicles are, in order from the eardrum to the inner ear (from superficial to deep): the
malleus The malleus, or hammer, is a hammer-shaped small bone or ossicle of the middle ear. It connects with the incus, and is attached to the inner surface of the eardrum. The word is Latin for 'hammer' or 'mallet'. It transmits the sound vibrations f ...
,
incus The ''incus'' (plural incudes) or anvil is a bone in the middle ear. The anvil-shaped small bone is one of three ossicles in the middle ear. The ''incus'' receives vibrations from the ''malleus'', to which it is connected laterally, and transmit ...
, and
stapes The ''stapes'' or stirrup is a bone in the middle ear of humans and other animals which is involved in the conduction of sound vibrations to the inner ear. This bone is connected to the oval window by its annular ligament, which allows the foo ...
, terms that in
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
are translated as "the hammer, anvil, and stirrup". * The
malleus The malleus, or hammer, is a hammer-shaped small bone or ossicle of the middle ear. It connects with the incus, and is attached to the inner surface of the eardrum. The word is Latin for 'hammer' or 'mallet'. It transmits the sound vibrations f ...
( la, "hammer") articulates with the incus through the
incudomalleolar joint The incudomalleolar joint (more correctly called incudomallear joint) or articulatio incudomallearis is a small synovial joint between the malleus (hammer) and the incus (anvil). The joint's function is to transfer vibrations between the ossic ...
and is attached to the
tympanic membrane 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 from the middle ear. Its function is to transmit sound from the a ...
( eardrum), from which vibrational sound pressure motion is passed. * The
incus The ''incus'' (plural incudes) or anvil is a bone in the middle ear. The anvil-shaped small bone is one of three ossicles in the middle ear. The ''incus'' receives vibrations from the ''malleus'', to which it is connected laterally, and transmit ...
( la, "anvil") is connected to both the other bones. * The
stapes The ''stapes'' or stirrup is a bone in the middle ear of humans and other animals which is involved in the conduction of sound vibrations to the inner ear. This bone is connected to the oval window by its annular ligament, which allows the foo ...
( la, "stirrup") articulates with the incus through the
incudostapedial joint Incudostapedial joint is a small, synovial ball-and-socket joint between the '' incus'' (anvil) and the ''stapes'' (stirrup). The joint's function is to transfer vibrations between the two ossicles. The incudostapedial joint lies between the lon ...
and is attached to the membrane of the '' fenestra ovalis'', the elliptical or
oval window The oval window (or ''fenestra vestibuli'' or ''fenestra ovalis'') is a membrane-covered opening from the middle ear to the cochlea of the inner ear. Vibrations that contact the tympanic membrane travel through the three ossicles and into the in ...
or opening between the middle ear and the vestibule of the inner ear. It is the smallest bone in the body.


Development

Studies have shown that ear bones in mammal embryos are attached to the
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
, which is part of the lower
jaw The jaw is any opposable articulated structure at the entrance of the mouth, typically used for grasping and manipulating food. The term ''jaws'' is also broadly applied to the whole of the structures constituting the vault of the mouth and serv ...
. These are ossified portions of cartilage—called
Meckel's cartilage In humans, the cartilaginous bar of the mandibular arch is formed by what are known as Meckel's cartilages (right and left) also known as Meckelian cartilages; above this the incus and malleus are developed. Meckel's cartilage arises from the fir ...
—that are attached to the jaw. As the embryo develops, the cartilage hardens to form bone. Later in development, the bone structure breaks loose from the jaw and migrates to the inner ear area. The structure is known as the middle ear, and is made up of the
stapes The ''stapes'' or stirrup is a bone in the middle ear of humans and other animals which is involved in the conduction of sound vibrations to the inner ear. This bone is connected to the oval window by its annular ligament, which allows the foo ...
,
incus The ''incus'' (plural incudes) or anvil is a bone in the middle ear. The anvil-shaped small bone is one of three ossicles in the middle ear. The ''incus'' receives vibrations from the ''malleus'', to which it is connected laterally, and transmit ...
,
malleus The malleus, or hammer, is a hammer-shaped small bone or ossicle of the middle ear. It connects with the incus, and is attached to the inner surface of the eardrum. The word is Latin for 'hammer' or 'mallet'. It transmits the sound vibrations f ...
, and
tympanic membrane 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 from the middle ear. Its function is to transmit sound from the a ...
. These correspond to the
columella Lucius Junius Moderatus Columella (; Arabic: , 4 – ) was a prominent writer on agriculture in the Roman Empire. His ' in twelve volumes has been completely preserved and forms an important source on Roman agriculture, together with the wo ...
, quadrate,
articular The articular bone is part of the lower jaw of most vertebrates, including most jawed fish, amphibians, birds and various kinds of reptiles, as well as ancestral mammals. Anatomy In most vertebrates, the articular bone is connected to two oth ...
, and angular structures in the amphibian, bird or reptile jaw.


Evolution


Function

As sound waves vibrate the tympanic membrane (eardrum), it in turn moves the nearest ossicle, the malleus, to which it is attached. The malleus then transmits the vibrations, via the incus, to the stapes, and so ultimately to the membrane of the fenestra ovalis (oval window), the opening to the vestibule of the inner ear. Sound traveling through the air is mostly reflected when it comes into contact with a liquid medium; only about 1/30 of the sound energy moving through the air would be transferred into the liquid. This is observed from the abrupt cessation of sound that occurs when the head is submerged underwater. This is because the relative incompressibility of a liquid presents resistance to the force of the sound waves traveling through the air. The ossicles give the eardrum a
mechanical advantage Mechanical advantage is a measure of the force amplification achieved by using a tool, mechanical device or machine system. The device trades off input forces against movement to obtain a desired amplification in the output force. The model for ...
via lever action and a reduction in the area of force distribution; the resulting vibrations are stronger but don't move as far. This allows more efficient coupling than if the sound waves were transmitted directly from the outer ear to the oval window. This reduction in the area of force application allows a large enough increase in
pressure Pressure (symbol: ''p'' or ''P'') is the force applied perpendicular to the surface of an object per unit area over which that force is distributed. Gauge pressure (also spelled ''gage'' pressure)The preferred spelling varies by country and e ...
to transfer most of the sound energy into the liquid. The increased pressure will compress the fluid found in the cochlea and transmit the stimulus. Thus, the lever action of the ossicles changes the vibrations so as to improve the transfer and reception of sound, and is a form of
impedance matching In electronics, impedance matching is the practice of designing or adjusting the input impedance or output impedance of an electrical device for a desired value. Often, the desired value is selected to maximize power transfer or minimize si ...
. However, the extent of the movements of the ossicles is controlled (and constricted) by two muscles attached to them (the ''
tensor tympani The tensor tympani is a muscle within the middle ear, located in the bony canal above the bony part of the auditory tube, and connects to the malleus bone. Its role is to dampen loud sounds, such as those produced from chewing, shouting, or th ...
'' and the '' stapedius''). It is believed that these muscles can contract to dampen the vibration of the ossicles, in order to protect the inner ear from excessively loud noise (theory 1) and that they give better frequency resolution at higher frequencies by reducing the transmission of low frequencies (theory 2) (see
acoustic reflex The acoustic reflex (also known as the stapedius reflex, stapedial reflex, auditory reflex, middle-ear-muscle reflex (MEM reflex, MEMR), attenuation reflex, cochleostapedial reflex or intra-aural reflex) is an involuntary muscle contraction that ...
). These muscles are more highly developed in
bat 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 ...
s and serve to block outgoing cries of the bats during echolocation (SONAR).


Clinical relevance

Occasionally the joints between the ossicles become rigid. One condition,
otosclerosis Otosclerosis is a condition of the middle ear where portions of the dense enchondral layer of the bony labyrinth remodel into one or more lesions of irregularly-laid spongy bone. As the lesions reach the stapes the bone is resorbed, then har ...
, results in the fusing of the stapes to the oval window. This reduces
hearing 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 audit ...
and may be treated surgically using a passive
middle ear implant A middle ear implant is a hearing device that is surgically implanted into the middle ear. They help people with conductive, sensorineural or mixed hearing loss to hear.   Middle ear implants work by improving the conduction of sound vibrations fr ...
.


History

There is some doubt as to the discoverers of the auditory ossicles and several anatomists from the early 16th century have the discovery attributed to them with the two earliest being
Alessandro Achillini Alessandro Achillini (''Latin'' Alexander Achillinus; 20 or 29 October 1463 (or possibly 1461)2 August 1512) was an Italian philosopher and physician. He is known for the anatomic studies that he was able to publish, made possible by a 13th-cent ...
and
Jacopo Berengario da Carpi Jacopo Berengario da Carpi (also known as Jacobus Berengarius Carpensis, Jacopo Barigazzi, Giacomo Berengario da Carpi or simply Carpus; c. 1460 – c. 1530) was an Italian physician. His book "''Isagoge breves''" published in 1522 made him the mo ...
. Several sources, including Eustachi and Casseri, attribute the discovery of the malleus and incus to the anatomist and philosopher Achillini. The first written description of the malleus and incus was by
Berengario da Carpi Jacopo Berengario da Carpi (also known as Jacobus Berengarius Carpensis, Jacopo Barigazzi, Giacomo Berengario da Carpi or simply Carpus; c. 1460 – c. 1530) was an Italian physician. His book "''Isagoge breves''" published in 1522 made him the mo ...
in his ''Commentaria super anatomia Mundini'' (1521), although he only briefly described two bones and noted their theoretical association with the transmission of sound.O'Malley, C.D. ''Andreas Vesalius of Brussels, 1514–1564.'' Berkeley: University of California Press, 1964. p. 120 Niccolo Massa's ''Liber introductorius anatomiae'' described the same bones in slightly more detail and likened them both to little hammers. A much more detailed description of the first two ossicles followed in
Andreas Vesalius Andreas Vesalius (Latinized from Andries van Wezel) () was a 16th-century anatomist, physician, and author of one of the most influential books on human anatomy, ''De Humani Corporis Fabrica Libri Septem'' (''On the fabric of the human body'' ' ...
' ''
De humani corporis fabrica ''De Humani Corporis Fabrica Libri Septem'' (Latin, lit. "On the fabric of the human body in seven books") is a set of books on human anatomy written by Andreas Vesalius (1514–1564) and published in 1543. It was a major advance in the history ...
'' in which he devoted a chapter to them. Vesalius was the first to compare the second element of the ossicles to an anvil although he offered the molar as an alternative comparison for its shape. The first published description of the stapes came in Pedro Jimeno's ''Dialogus de re medica'' (1549) although it had been previously described in public lectures by Giovanni Filippo Ingrassia at the
University of Naples The University of Naples Federico II ( it, Università degli Studi di Napoli Federico II) is a public university in Naples, Italy. Founded in 1224, it is the oldest public non-sectarian university in the world, and is now organized into 26 depar ...
as early as 1546. The term ''ossicle'' derives from , a diminutive of "bone" ( la, os; genitive ). The malleus gets its name from Latin ''malleus'', meaning "hammer", the incus gets its name from Latin ''incus'' meaning "anvil" from incudere meaning "to forge with a hammer", and the stapes gets its name from Modern Latin "stirrup", probably an alteration of Late Latin ''stapia'' related to stare "to stand" and pedem, an accusative of pes "foot", so called because the bone is shaped like a stirrup – this was an invented Modern Latin word for "stirrup", for which there was no classical Latin word, as the ancients did not use stirrups.


See also

* * *


References


External links


The middle ear and the ossicles
{{Authority control Auditory system Bones of the head and neck