The utricle and
saccule
The saccule is a bed of sensory cells in the inner ear. It translates head movements into neural impulses for the brain to interpret. The saccule detects linear accelerations and head tilts in the vertical plane. When the head moves verticall ...
are the two
otolith
An otolith ( grc-gre, ὠτο-, ' ear + , ', a stone), also called statoconium or otoconium or statolith, is a calcium carbonate structure in the saccule or utricle of the inner ear, specifically in the vestibular system of vertebrates. The sa ...
organs in the
vertebrate
Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
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 ...
. They are part of the balancing system (
membranous labyrinth
The membranous labyrinth is a collection of fluid filled tubes and chambers which contain the receptors for the senses of equilibrium and hearing. It is lodged within the bony labyrinth in the inner ear and has the same general form; it is, howeve ...
) in the vestibule of the
bony labyrinth
The bony labyrinth (also osseous labyrinth or otic capsule) is the rigid, bony outer wall of the inner ear in the temporal bone. It consists of three parts: the vestibule, semicircular canals, and cochlea. These are cavities hollowed out of the s ...
(small oval chamber). They use small stones and a viscous fluid to stimulate hair cells to detect motion and orientation. The utricle detects linear accelerations and head-tilts in the horizontal plane. The word utricle comes .
Structure
The utricle is larger than the saccule and is of an oblong form, compressed transversely, and occupies the upper and back part of the
vestibule, lying in contact with the
recessus ellipticus and the part below it.
Macula
The macula of utricle (macula acustica utriculi) is a small (2 by 3 mm) thickening lying horizontally on the floor of the utricle where the epithelium contains vestibular hair cells that allow a person to perceive changes in latitudinal acceleration as well as the effects of gravity; it receives the utricular filaments of the
acoustic nerve
The cochlear nerve (also auditory nerve or acoustic nerve) is one of two parts of the vestibulocochlear nerve, a cranial nerve present in amniotes, the other part being the vestibular nerve. The cochlear nerve carries auditory sensory information ...
.
The hair cells are mechanoreceptors which have 40 to 70 stereocilia and only one true cilium called a
kinocilium
A kinocilium is a special type of cilium on the apex of hair cells located in the sensory epithelium of the vertebrate inner ear.
Anatomy in humans
Kinocilia are found on the apical surface of hair cells and are involved in both the morphogenesis ...
. The kinocilium is the only sensory aspect of the hair cell and is what causes hair cell polarization. The tips of these stereocilia and kinocilium are embedded in a gelatinous layer, which together with the
statoconia
An otolith ( grc-gre, ὠτο-, ' ear + , ', a stone), also called statoconium or otoconium or statolith, is a calcium carbonate structure in the saccule or utricle of the inner ear, specifically in the vestibular system of vertebrates. The sa ...
form the
otolithic membrane. This membrane is weighted with calcium carbonate-protein granules called otoliths. The otolithic membrane adds weight to the tops of the hair cells and increases their inertia. The addition in weight and inertia is vital to the utricle's ability to detect linear acceleration, as described below, and to determine the orientation of the head. When the head is tilted such that gravity pulls on the statoconia, the gelatinous layer is pulled in the same direction also, causing the sensory hairs to bend. Labyrinthine activity responsible for the
nystagmus
Nystagmus is a condition of involuntary (or voluntary, in some cases) eye movement. Infants can be born with it but more commonly acquire it in infancy or later in life. In many cases it may result in reduced or limited vision. Due to the invol ...
induced by off-vertical axis rotation arises in the otolith organs and couples to the oculomotor system through the velocity storage mechanism.
Microanatomy
The cavity of the utricle communicates behind with the
semicircular ducts
In mathematics (and more specifically geometry), a semicircle is a one-dimensional locus of points that forms half of a circle. The full arc of a semicircle always measures 180° (equivalently, radians, or a half-turn). It has only one line o ...
by five orifices.
The ductus utriculosaccularis comes off of the anterior wall of the utricle and opens into the
ductus endolymphaticus
From the posterior wall of the saccule a canal, the endolymphatic duct, is given off; this duct is joined by the ductus utriculosaccularis, and then passes along the aquaeductus vestibuli and ends in a blind pouch (endolymphatic sac) on the poste ...
.
Function
The utricle contains mechanoreceptors called hair cells that distinguish between degrees of tilting of the head, thanks to their apical
stereocilia
Stereocilia (or stereovilli or villi) are non-motile apical cell modifications. They are distinct from cilia and microvilli, but are closely related to microvilli. They form single "finger-like" projections that may be branched, with normal cel ...
set-up. These are covered by
otolith
An otolith ( grc-gre, ὠτο-, ' ear + , ', a stone), also called statoconium or otoconium or statolith, is a calcium carbonate structure in the saccule or utricle of the inner ear, specifically in the vestibular system of vertebrates. The sa ...
s which, due to gravity, pull on the stereocilia and tilt them. Depending on whether the tilt is in the direction of the
kinocilium
A kinocilium is a special type of cilium on the apex of hair cells located in the sensory epithelium of the vertebrate inner ear.
Anatomy in humans
Kinocilia are found on the apical surface of hair cells and are involved in both the morphogenesis ...
or not, the resulting hair cell polarisation is excitatory (depolarising) or inhibitory (hyperpolarisation), respectively. Any orientation of the head causes a combination of stimulation to the utricles and saccules of the two ears. The brain interprets head orientation by comparing these inputs to each other and to other input from the eyes and stretch receptors in the neck, thereby detecting whether only the head is tilted or the entire body is tipping. The inertia of the otolithic membranes is especially important in detecting linear acceleration. Suppose you are sitting in a car at a stoplight and then begin to move. The otolithic membrane of the macula utriculi briefly lags behind the rest of the tissues, bends the stereocilia backward, and stimulates the cells. When you stop at the next light, the macula stops but the otolithic membrane keeps going for a moment, bending the stereocilia forward. The hair cells convert this pattern of stimulation to nerve signals, and the brain is thus advised of changes in your linear velocity.
[Saladin, Kenneth S. Anatomy & Physiology: the Unity of Form and Function. Dubuque: McGraw-Hill, 2010. Print.] This signal to the vestibular nerve (which takes it to the brainstem) does not adapt with time. The effect of this is that, for example, an individual lying down to sleep will continue to detect that they are lying down hours later when they awaken.
Unbent and at rest hairs in the macula have a base rate of depolarization of 90-100 action potentials a second. The brain suppresses this, and we ignore it and know that our body is stabilized. If the head moves or the body accelerates or decelerates, then bending occurs. Depending on the direction of bending, the hair cells will either be excited or inhibited resulting in either an increase or decrease in firing frequency of the hair cells.
The macula is also sensitive to linear acceleration as the inertia possessed by the statoconia can also shift the gelatinous layer during increases and decreases in linear velocity.
See also
References
External links
Diagram at ipfw.edu
{{Authority control
Ear
Vestibular system