The middle ear is the portion of the ear medial to the

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 sound from the ...

, and distal to the

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 ...

of the cochlea (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 th ...

). The mammalian middle ear contains three

ossicles The ossicles (also called auditory ossicles) are three bones in either middle ear that are among the smallest bones in the human body. They serve to transmit sounds from the air to the fluid-filled labyrinth (cochlea). The absence of the auditor ...

, which transfer the vibrations of the eardrum into waves in the fluid and membranes of the

. The hollow space of the middle ear is also known as the

tympanic cavity The tympanic cavity is a small cavity surrounding the bones of the middle ear. Within it sit the ossicles, three small bones that transmit vibrations used in the detection of sound. Structure On its lateral surface, it abuts the external auditor ...

and is surrounded by the tympanic part of the temporal bone. The

auditory tube Auditory means of or relating to the process of hearing: * Auditory system, the neurological structures and pathways of sound perception ** Auditory bulla, part of auditory system found in mammals other than primates ** Auditory nerve, also known ...

(also known as the Eustachian tube or the pharyngotympanic tube) joins the tympanic cavity with the nasal cavity (

nasopharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its struct ...

), allowing pressure to equalize between the middle ear and throat. The primary function of the middle ear is to efficiently transfer acoustic energy from

compression wave Longitudinal waves are waves in which the vibration of the medium is parallel ("along") to the direction the wave travels and displacement of the medium is in the same (or opposite) direction of the wave propagation. Mechanical longitudinal waves ...

s in air to fluid–membrane waves within the

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 o ...

Structure

Ossicles

The middle ear contains three tiny bones known as the

: '' malleus'', '' incus'', 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 ...

''. The ossicles were given their Latin names for their distinctive shapes; they are also referred to as the ''hammer'', ''anvil'', and ''stirrup'', respectively. The ossicles directly couple sound energy from the eardrum to the

of the cochlea. While the stapes is present in all

tetrapods Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct therapsi ...

, the malleus and incus evolved from lower and upper jaw bones present in reptiles. The ossicles are classically supposed to mechanically convert the vibrations of the

into amplified pressure waves in the fluid of the

(or

), with a lever arm factor of 1.3. Since the effective vibratory area of the eardrum is about 14 fold larger than that of the oval window, the sound pressure is concentrated, leading to a pressure gain of at least 18.1. The eardrum is merged to the malleus, which connects to the incus, which in turn connects to the stapes. Vibrations of the stapes footplate introduce pressure waves in the

. There is a steadily increasing body of evidence that shows that the lever arm ratio is actually variable, depending on frequency. Between 0.1 and 1 kHz it is approximately 2, it then rises to around 5 at 2 kHz and then falls off steadily above this frequency. The measurement of this lever arm ratio is also somewhat complicated by the fact that the ratio is generally given in relation to the tip of the malleus (also known as the umbo) and the level of the middle of the stapes. The eardrum is actually attached to the malleus handle over about a 0.5 cm distance. In addition, the eardrum itself moves in a very chaotic fashion at frequencies >3 kHz. The linear attachment of the eardrum to the malleus actually smooths out this chaotic motion and allows the ear to respond linearly over a wider frequency range than a point attachment. The auditory ossicles can also reduce sound pressure (the inner ear is very sensitive to overstimulation), by uncoupling each other through particular muscles. The middle ear efficiency peaks at a frequency of around 1 kHz. The combined

transfer function In engineering, a transfer function (also known as system function or network function) of a system, sub-system, or component is a mathematical function that theoretically models the system's output for each possible input. They are widely used ...

of the outer ear and middle ear gives humans a peak sensitivity to frequencies between 1 kHz and 3 kHz.

Muscles

The movement of the ossicles may be stiffened by two muscles. The

stapedius muscle 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 strirrup bone of the middle ear. Structure The stapedius emerges from ...

, the smallest skeletal muscle in the body, connects to the stapes and is controlled by the

facial nerve The facial nerve, also known as the seventh cranial nerve, cranial nerve VII, or simply CN VII, is a cranial nerve that emerges from the pons of the brainstem, controls the muscles of facial expression, and functions in the conveyance of taste ...

; the

tensor tympani muscle 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 thu ...

is attached to the upper end of the medial surface of the handle of malleus and is under the control of the

medial pterygoid nerve The medial pterygoid nerve (or internal pterygoid nerve) is a nerve of the head. It is a branch of the mandibular nerve (CN V3). It supplies the medial pterygoid muscle, the tensor veli palatini muscle, and the tensor tympani muscle. Structure ...

which is a branch of the

mandibular nerve In neuroanatomy, the mandibular nerve (V) is the largest of the three divisions of the trigeminal nerve, the fifth cranial nerve (CN V). Unlike the other divisions of the trigeminal nerve ( ophthalmic nerve, maxillary nerve) which contain only ...

of the

trigeminal nerve In neuroanatomy, the trigeminal nerve ( lit. ''triplet'' nerve), also known as the fifth cranial nerve, cranial nerve V, or simply CN V, is a cranial nerve responsible for sensation in the face and motor functions such as biting and che ...

. These muscles contract in response to loud sounds, thereby reducing the transmission of sound to the inner ear. This is called the acoustic reflex.

Nerves

Of surgical importance are two branches of the

that also pass through the middle ear space. These are the horizontal portion of the facial nerve and the

chorda tympani The chorda tympani is a branch of the facial nerve that originates from the taste buds in the front of the tongue, runs through the middle ear, and carries taste messages to the brain. It joins the facial nerve (cranial nerve VII) inside the fa ...

. Damage to the horizontal branch during ear surgery can lead to paralysis of the face (same side of the face as the ear). The chorda tympani is the branch of the facial nerve that carries taste from the

ipsilateral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...

half (same side) of the tongue.

Function

Sound transfer

Ordinarily, when sound waves in air strike liquid, most of the energy is reflected off the surface of the liquid. The middle ear allows the

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 ...

of sound traveling in air to acoustic waves traveling in a system of fluids and membranes in the inner ear. This system should not be confused, however, with the propagation of sound as compression waves in liquid. The middle ear couples sound from air to the fluid via the

, using the principle of "mechanical advantage" in the form of the "hydraulic principle" and the "lever principle". The vibratory portion of the tympanic membrane (eardrum) is many times the surface area of the footplate of the

(the third ossicular bone which attaches to the oval window); furthermore, the shape of the articulated ossicular chain is a complex

lever A lever is a simple machine consisting of a beam or rigid rod pivoted at a fixed hinge, or '' fulcrum''. A lever is a rigid body capable of rotating on a point on itself. On the basis of the locations of fulcrum, load and effort, the lever is d ...

, the long arm being the long process of the malleus, the fulcrum being the body of the incus, and the short arm being the lenticular process of the incus. The collected pressure of sound vibration that strikes the tympanic membrane is therefore concentrated down to this much smaller area of the footplate, increasing the force but reducing the velocity and displacement, and thereby coupling the acoustic energy. The middle ear is able to dampen sound conduction substantially when faced with very loud sound, by noise-induced reflex contraction of the middle-ear muscles.

Clinical significance

The middle ear is hollow. In a high-altitude environment or on diving into water, there will be a pressure difference between the middle ear and the outside environment. This pressure will pose a risk of bursting or otherwise damaging the tympanum if it is not relieved. If middle ear pressure remains low, the eardrum may become retracted into the middle ear. One of the functions of the

Eustachian tube In anatomy, the Eustachian tube, also known as the auditory tube or pharyngotympanic tube, is a tube that links the nasopharynx to the middle ear, of which it is also a part. In adult humans, the Eustachian tube is approximately long and in d ...

s that connect the middle ear to the

is to help keep middle ear pressure the same as air pressure. The Eustachian tubes are normally pinched off at the nose end, to prevent being clogged with

mucus Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It ...

, but they may be opened by lowering and protruding the jaw; this is why yawning or chewing helps relieve the pressure felt in the ears when on board an aircraft.

Otitis media Otitis media is a group of inflammatory diseases of the middle ear. One of the two main types is acute otitis media (AOM), an infection of rapid onset that usually presents with ear pain. In young children this may result in pulling at the ear, ...

is an inflammation of the middle ear.

Injuries

The middle ear is well protected from most minor external injuries by its internal location, but is vulnerable to pressure injury (

barotrauma Barotrauma is physical damage to body tissues caused by a difference in pressure between a gas space inside, or contact with, the body and the surrounding gas or liquid. The initial damage is usually due to over-stretching the tissues in tensi ...

Infections

Recent findings indicate that the middle ear mucosa could be subjected to

human papillomavirus Human papillomavirus infection (HPV infection) is caused by a DNA virus from the '' Papillomaviridae'' family. Many HPV infections cause no symptoms and 90% resolve spontaneously within two years. In some cases, an HPV infection persists and r ...

infection. Indeed, DNAs belonging to oncogenic HPVs, i.e., HPV16 and HPV18, have been detected in normal middle ear specimens, thereby indicating that the normal middle ear mucosa could potentially be a target tissue for HPV infection.

Other animals

The middle ear of

tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct t ...

s is analogous with the spiracle of fishes, an opening from the

pharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its st ...

to the side of the head in front of the main gill slits. In fish embryos, the spiracle forms as a pouch in the pharynx, which grows outward and breaches the skin to form an opening; in most tetrapods, this breach is never quite completed, and the final vestige of tissue separating it from the outside world becomes the eardrum. The inner part of the spiracle, still connected to the pharynx, forms the eustachian tube. In

reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates ( lizards and snakes) and rhynchocephalia ...

bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...

s, and early fossil tetrapods, there is a single auditory ossicle, 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 ...

which is homologous with the stapes, or "stirrup" of mammals. This is connected indirectly with the eardrum via a mostly cartilaginous extracolumella and medially to the inner-ear spaces via a widened footplate in the fenestra ovalis. The columella is an evolutionary derivative of the bone known as the hyomandibula in fish ancestors, a bone that supported the skull and braincase. The structure of the middle ear in living

amphibian Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbo ...

s varies considerably and is often degenerate. In most frogs and toads, it is similar to that of reptiles, but in other amphibians, the middle ear cavity is often absent. In these cases, the stapes either is also missing or, in the absence of an eardrum, connects to the quadrate bone in the skull, although, it is presumed, it still has some ability to transmit vibrations to the inner ear. In many amphibians, there is also a second auditory ossicle, the ''operculum'' (not to be confused with the structure of the same name in fishes). This is a flat, plate-like bone, overlying the fenestra ovalis, and connecting it either to the stapes or, via a special muscle, to the

scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eith ...

. It is not found in any other vertebrates. Mammals are unique in having evolved a three-ossicle middle-ear independently of the various single-ossicle middle ears of other land vertebrates, all during the Triassic period of geological history. Functionally, the mammalian middle ear is very similar to the single-ossicle ear of non-mammals, except that it responds to sounds of higher frequency, because these are better taken up by the inner ear (which also responds to higher frequencies than those of non-mammals). The malleus, or "hammer", evolved from the

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 ...

bone of the lower jaw, and the incus, or "anvil", from the quadrate. In other vertebrates, these bones form the primary jaw joint, but the expansion of 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 ...

bone in mammals led to the evolution of an entirely new jaw joint, freeing up the old joint to become part of the ear. For a period of time, both jaw joints existed together, one medially and one laterally. The evolutionary process leading to a three-ossicle middle ear was thus an "accidental" byproduct of the simultaneous evolution of the new, secondary jaw joint. In many mammals, the middle ear also becomes protected within a cavity, the

auditory bulla The tympanic part of the temporal bone is a curved plate of bone lying below the squamous part of the temporal bone, in front of the mastoid process, and surrounding the external part of the ear canal. It originates as a separate bone (tympanic ...

, not found in other vertebrates. A bulla evolved late in time and independently numerous times in different mammalian clades, and it can be surrounded by membranes, cartilage or bone. The bulla in humans is part of the

temporal bone The temporal bones are situated at the sides and base of the skull, and lateral to the temporal lobes of the cerebral cortex. The temporal bones are overlaid by the sides of the head known as the temples, and house the structures of the ears. ...

Additional images

File:Gray919.png

References

External links

Promenade Around the Cochlea - Middle ear at iurc.montp.inserm.fr
{{Authority control Auditory system Ear