/balance system. Arrows show the direction of information relay. This overview also explains acceleration as its processes are interconnected with balance.
Mechanical There are five
sensory organs innervated by the
vestibular nerve; three
semicircular canals (
Horizontal SCC,
Superior SCC,
Posterior SCC) and two
otolith organs (
saccule and
utricle). Each semicircular canal (SSC) is a thin tube that doubles in thickness briefly at a point called
osseous ampullae. At their center-base, each contains an
ampullary cupula. The cupula is a gelatin bulb connected to the
stereocilia of hair cells, affected by the relative movement of the
endolymph it is bathed in. Since the cupula is part of the
bony labyrinth, it rotates along with actual head movement, and by itself without the endolymph, it cannot be stimulated and therefore, could not detect movement. Endolymph follows the rotation of the canal; however, due to
inertia its movement initially lags behind that of the bony labyrinth. The delayed movement of the endolymph bends and activates the cupula. When the cupula bends, the connected stereocilia bend along with it, activating chemical reactions in the hair cells surrounding
crista ampullaris and eventually create
action potentials carried by the vestibular nerve signaling to the body that it has moved in space. After any extended rotation, the endolymph catches up to the canal and the cupula returns to its upright position and resets. When extended rotation ceases, however, endolymph continues, (due to inertia) which bends and activates the cupula once again to signal a change in movement. Pilots doing long banked turns begin to feel upright (no longer turning) as endolymph matches canal rotation; once the pilot exits the turn the cupula is once again stimulated, causing the feeling of turning the other way, rather than flying straight and level. The horizontal SCC handles head rotations about a vertical axis (e.g. looking side to side), the superior SCC handles head movement about a lateral axis (e.g. head to shoulder), and the posterior SCC handles head rotation about a
rostral-caudal axis (e.g. nodding). SCC sends adaptive signals, unlike the two otolith organs, the saccule and utricle, whose signals do not adapt over time. A shift in the
otolithic membrane that stimulates the cilia is considered the state of the body until the cilia are once again stimulated. For example, lying down stimulates cilia and standing up stimulates cilia, however, for the time spent lying the signal that you are lying remains active, even though the membrane resets.
Otolithic organs have a thick, heavy gelatin membrane that, due to inertia (like endolymph), lags behind and continues ahead past the
macula it overlays, bending and activating the contained cilia.
Utricle responds to linear accelerations and head-tilts in the horizontal plane (head to shoulder), whereas
saccule responds to linear accelerations and head-tilts in the vertical plane (up and down). Otolithic organs update the brain on the head-location when not moving; SCC update during movement.
Kinocilium are the longest stereocilia and are positioned (one per 40-70 regular cilia) at the end of the bundle. If stereocilia go towards kinocilium,
depolarization occurs, causing more
neurotransmitters, and more vestibular nerve firings, as compared to when stereocilia tilt away from kinocilium (
hyperpolarization, less neurotransmitter, less firing).
Neural First order
vestibular nuclei (VN) project to
lateral vestibular nucleus (IVN),
medial vestibular nucleus (MVN), and
superior vestibular nucleus (SVN). The
inferior cerebellar peduncle is the largest center through which balance information passes. It is the area of integration between
proprioceptive, and vestibular inputs, to aid in unconscious maintenance of balance and posture. The
inferior olivary nucleus aids in complex
motor tasks by encoding coordinating timing sensory information; this is decoded and acted upon in the
cerebellum. The
cerebellar vermis has three main parts. The
vestibulocerebellum regulates eye movements by the integration of visual info provided by the
superior colliculus and balance information. The
spinocerebellum integrates visual, auditory, proprioceptive, and balance information to act out body and limb movements. It receives input from the
trigeminal nerve, dorsal column (of the
spinal cord),
midbrain,
thalamus,
reticular formation and vestibular nuclei (
medulla) outputs. Lastly, the
cerebrocerebellum plans, times, and initiates movement after evaluating sensory input from, primarily, motor cortex areas, via
pons and cerebellar
dentate nucleus. It outputs to the thalamus,
motor cortex areas, and
red nucleus. The
flocculonodular lobe is a cerebellar lobe that helps maintain body equilibrium by modifying
muscle tone (the continuous and passive muscle contractions). MVN and IVN are in the medulla, LVN and SVN are smaller and in pons. SVN, MVN, and IVN ascend within the
medial longitudinal fasciculus. LVN descend the spinal cord within the
lateral vestibulospinal tract and ends at the
sacrum. MVN also descend the spinal cord, within the
medial vestibulospinal tract, ending at
lumbar 1. The
thalamic reticular nucleus distributes information to various other thalamic nuclei, regulating the flow of information. It is speculatively able to stop signals, ending transmission of unimportant info. The thalamus relays info between pons (cerebellum link), motor cortices, and
insula. The insula is also heavily connected to motor cortices; the insula is likely where balance is likely brought into perception. The
oculomotor nuclear complex refers to fibers going to
tegmentum (eye movement), red nucleus (
gait (natural limb movement)),
substantia nigra (reward), and
cerebral peduncle (motor relay). Nucleus of Cajal are one of the named oculomotor nuclei, they are involved in eye movements and reflex gaze coordination. The
abducens nerve solely innervates the
lateral rectus muscle of the eye, moving the eye with the
trochlear nerve. The trochlear solely innervates the
superior oblique muscle of the eye. Together, trochlear and abducens contract and relax to simultaneously direct the pupil towards an angle and depress the globe on the opposite side of the eye (e.g. looking down directs the pupil down and depresses (towards the brain) the top of the globe). The pupil is not only directed, but often rotated, by these muscles. (See
visual system) The thalamus and superior colliculus are connected via the
lateral geniculate nucleus. The superior colliculus (SC) is the
topographical map for balance and quick orienting movements with primarily visual inputs. SC integrates multiple senses. and
saccule organs that are responsible for detecting linear acceleration, or movement in a straight line. ==Other animals==