The ossicles (also called auditory ossicles) are three

irregular bone The irregular bones are bones which, from their peculiar form, cannot be grouped as long, short, flat or sesamoid bones. Irregular bones serve various purposes in the body, such as protection of nervous tissue (such as the vertebrae protect the ...

s in the

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 (malleus, incus, and stapes), which transfer the vibrations ...

human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...

s and other

mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...

s, and are among the smallest

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 the human body. Although the term "ossicle" literally means "tiny bone" (from

Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...

''ossiculum'') and may refer to any small bone throughout the body, it typically refers specifically 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 vibra ...

incus The ''incus'' (: incudes) or anvil in the ear is one of three small bones (ossicles) in the middle ear. The incus receives vibrations from the malleus, to which it is connected laterally, and transmits these to the stapes medially. The incus i ...

and

stapes The ''stapes'' or stirrup is a bone in the middle ear of humans and other tetrapods 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 f ...

("hammer, anvil, and stirrup") of the middle ear. The auditory ossicles serve as a kinematic chain to transmit and amplify (

intensify ''Intensify'' is the second studio album by English electronic music duo Way Out West, released on 20 August 2001 by Distinct'ive Records. The album was marked by a move from the defunct Deconstruction Records label to Distinctive, and a chang ...

)

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'' by the br ...

vibrations collected from the air by the

ear drum 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 changes in pressur ...

to the fluid-filled

labyrinth In Greek mythology, the Labyrinth () is 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 the h ...

(

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 (cochlea), modiolus. A core component of the cochlea is the organ of Cort ...

). The absence or pathology of the auditory ossicles would constitute a moderate-to-severe

conductive hearing loss Conductive hearing loss (CHL) is a type of hearing impairment that occurs when sound waves are unable to efficiently travel through the outer ear, tympanic membrane (eardrum), or middle ear structures such as the ossicles. This blockage or dysfun ...

Structure

The ossicles are, in order from the eardrum to the inner ear (from superficial to deep): the

, and

, terms that in

are translated as "the

hammer A hammer is a tool, most often a hand tool, consisting of a weighted "head" fixed to a long handle that is swung to deliver an impact to a small area of an object. This can be, for example, to drive nail (fastener), nails into wood, to sh ...

anvil An anvil is a metalworking tool consisting of a large block of metal (usually Forging, forged or Steel casting, cast steel), with a flattened top surface, upon which another object is struck (or "worked"). Anvils are massive because the hi ...

, and

stirrup A stirrup is a light frame or ring that holds the foot of a rider, attached to the saddle by a strap, often called a ''stirrup leather''. Stirrups are usually paired and are used to aid in mounting and as a support while using a riding animal ...

". * The

() articulates with the incus through the

incudomalleolar joint The incudomalleolar joint (also called incudomallear joint, or malleoincudal joint) or articulatio incudomallearis is a small saddle-type synovial joint between the malleus (hammer) and the incus (anvil). The joint's function is to transfer vib ...

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 changes in pressur ...

(

eardrum 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 changes in pres ...

), from which vibrational sound pressure motion is passed. * The

() is connected to both the other bones. * The

() articulates with the incus through the

incudostapedial joint The 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 ...

and is attached to the membrane of the ''

fenestra ovalis The oval window (or ''fenestra vestibuli'' or ''fenestra ovalis'') is a connective tissue membrane-covered opening from the middle ear to the cochlea of the inner ear. Vibrations that contact the tympanic membrane travel through the three ossicl ...

'', the elliptical or

oval window The oval window (or ''fenestra vestibuli'' or ''fenestra ovalis'') is a connective tissue membrane-covered opening from the middle ear to the cochlea of the inner ear. Vibrations that contact the tympanic membrane travel through the three ossicl ...

or opening between the middle ear and the vestibule of the

inner ear The inner ear (internal ear, auris interna) is the innermost part of the vertebrate ear. In vertebrates, the inner ear is mainly responsible for sound detection and balance. In mammals, it consists of the bony labyrinth, a hollow cavity in the ...

. It is the smallest bone in the body.

Development

Studies have shown that ear bones in mammal embryos are attached to the

dentary In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone ...

, which is part of the lower

jaw The jaws are a pair of opposable articulated structures 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 ...

. These are ossified portions of

cartilage Cartilage is a resilient and smooth type of connective tissue. Semi-transparent and non-porous, it is usually covered by a tough and fibrous membrane called perichondrium. In tetrapods, it covers and protects the ends of long bones at the joints ...

—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 which the incus and malleus are located. 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

, and

. These correspond to the

columella Lucius Junius Moderatus Columella (, Arabic: ) was a prominent Roman writer on agriculture in the Roman Empire. His in twelve volumes has been completely preserved and forms an important source on Roman agriculture and ancient Roman cuisin ...

, 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 o ...

, 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 eve ...

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 electrical engineering, 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 ...

. 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 thun ...

'' and the ''

stapedius The stapedius is the smallest skeletal muscle in the human body. At just over one millimeter in length, its purpose is to stabilize the smallest bone in the body, the stapes or stirrup bone of the middle ear. Structure The stapedius emerges from ...

''). 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 flying mammals of the order Chiroptera (). 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 birds, flying with their very long spread-out ...

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, middle and inner ear where portions of the dense enchondral layer of the bony labyrinth Tissue remodeling, remodel into one or more lesions of irregularly-laid spongy bone. As the lesions reach the s ...

, 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 auditory sci ...

and may be treated surgically using a passive middle ear implant.

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 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 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 Andries van Wezel (31 December 1514 – 15 October 1564), latinized as Andreas Vesalius (), was an anatomist and physician who wrote '' De Humani Corporis Fabrica Libri Septem'' (''On the fabric of the human body'' ''in seven books''), which is ...

' ''

De humani corporis fabrica ''De Humani Corporis Fabrica Libri Septem'' (Latin, "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 of a ...

'' 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 (; , ) is a public university, public research university in Naples, Campania, Italy. Established in 1224 and named after its founder, Frederick II, Holy Roman Emperor, Frederick II, it is the oldest public, s ...

as early as 1546. The term ''ossicle'' derives from , a

diminutive A diminutive is a word obtained by modifying a root word to convey a slighter degree of its root meaning, either to convey the smallness of the object or quality named, or to convey a sense of intimacy or endearment, and sometimes to belittle s ...

of "bone" (;

genitive In grammar, the genitive case ( abbreviated ) is the grammatical case that marks a word, usually a noun, as modifying another word, also usually a noun—thus indicating an attributive relationship of one noun to the other noun. A genitive can ...

). 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.

References

External links

The middle ear and the ossicles
{{Authority control Auditory system Bones of the head and neck Otorhinolaryngology Otology Human head and neck Ear Middle ear

Structure

Development

Evolution

Function

Clinical relevance

History

See also

References

External links