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Vestibular sensitivity
Francisco E. Olucha-Bordonau
Universitat de València

Content

Vestibular sensitivity .............................................................................................................. 1

The vestibular organ .......................................................................................................... 1

The vestibular complex ...................................................................................................... 1

Proprioceptive pathways .................................................................................................... 2

Proprioception and vestibular representation in the cortex .................................................. 4

Vestibular sensitivity
The statoacoustic nerve brings together the sensibilities of the cochlear and vestibular organs,
but these informations do not mix at any time. The vestibular nerve passes between the inferior
cerebellar peduncle and the trigeminospinal tract and penetrate the vestibular nuclear complex

The vestibular organ

The vestibular organ corresponds to the lateral aspect of the laberinth containing the
semicircular ducts, the utriculum and the saculum. The sensory epithelium of the utricle and
saculum are called maculae. The macula of the utricle lies on the floor of the utricle parallel to
the base of the skull, thus nearly horizontal. In contrast, the macula of saculum stands vertically
on the medial side. The main component of the maculae’s epithelium are ciliated cells that lies
on a base containing the terminal ending of the ganglion neurons. On the apical part the ciliae
are ambedded on a meambrane and the endolymph contain cristaline deposits, named otolits,
that may fall on the membraine depending on the position of the body like in a snow ball. Thus
the combination of the four planes (utricle and saculum in each side) sends the signal about the
position of the head in the space.

In the semicircular duct epithelium sensitive ampulla is located on a ridge up a plane that is
perpendicular to the axis of the duct and making relief in its lumen. The epithelium is composed
of the crest cells and ciliated sensitive cells and is covered by a gelatinous dome containing the
cilia. The head movements cause the opposite movement of endolymph inside the semicircular
canal what causes the activation of ciliated cells. On the basal part of the ciliated cells there are
contacts with the terminal processes of the vestibular ganglion cells

Fibers coming from the vestibular sensitive organs arrive to the vestibularganglion (of Scarpa).
Actually, this ganglion consists of two parts, an upper one and a lower one. The upper
vestibular ganglion contains ganglion cells that collect information from the ampullae of anterior
and lateral ducts and the utricle. The bottom vestibular ganglion contains cells that collect
information from the saculum and the posterior semicircular duct.

The axons of ganglion cells in the vestibular ganglion and spiral ganglion join to form the joint
VIII cranial pair.

The vestibular complex

The vestibular complex is composed of four nuclei: lateral, superior, inferior and medial.
Superior, lateral and medial nuclei extend rostrocaudally in the same length as the medial
vestibular nucleus. The axons of vestibular ganglion cells are distributed on all four nuclei. On
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entering the complex are divided into T with a short ascending branch and a long descending
one. The crests of the semicircular ducts are projected primarily in the superior vestibular
nucleus and in the rostral part of the medial vestibular nuclei. The utricle and saculum are
projected primarily in the ventral part of the lateral and inferior vestibular nuclei.

Some ganglionar vestibular fibers passed directly to the vestibulo-cerebellum without relaying in
the vestibular nuclei through the juxtarestiform body.

In addition to the output of the vestibular nucleis. These nuclei display a system of intrinsic
ipsilateral and contralateral intrinsic connections. Also the vestibular system receive projections
from the accessory visual system which correlates vision with movement. This system is mainly
composed of pretectal centers like the nucleus of the optic tract. Research about the sensitivity
of neurons in the nucleus of the optic tract revels that these neurons are sensitive to wide field
and movements of 0.1-30 degrees/sec what fits to a sensitivity to automevement instead of
movement a particular object in the sapace. The vestibular nuclei also receive projections from
several pontine nuclei, the pedunculo-pontine complex the inferior olive and the nucleus
prepositus of the hipoglossum. All these nuclei together with the pretectal accessory visual
system provides an optional spatial map that is also configured in the vestibular nuclei.

The vestibular nuclei originate three kinds of connections to the cerebellum, the spinal cord and
the oculomotor centers. The vestibular projections to the cerebellum arises from the medial and
inferior nuclei and courses through the juxtarestiformis body.

The lateral vestibular nucleus originates the lateral vestibulo-spinal tract which courses through
the ipsilateral spinal cord. The projection covers all levels of the spinal cord and targets mainly
the layers VII and VIII in the anterior horn. This nucleus is under the control of the cerebellum
which makes possible mopre accurate movements. The medial vestibular nucleus medial
vestibulo-spinal tract, but only reaches the upper thoracic and cervical levels. The vestibular
influence on the spinal cord is facilitating spinal motor neurons firing to activate muscles. In
addition, the vestibular pathway over the cervical levels provides a stable base for any action of
the muscles of the eye movements.

Finally, the medial and superior vestibular nuclei regulate largely coordinated movements of the
head and eyes as they provide a large amount of fibers to the medial longitudinal fascicle. This
tract appears bellow the ventral part of the aqueduct and connects the ocular motor nuclei
between them and with the accessory nucleus which moves the trapezium and
sternoclidomastoideus. Thus, the movements of the neck and eyes are coordinated by the
information obtained from the angular acceleration of the head detected by the semicircular
ducts of the ampulla.

We should also mention that part of the vestibular fibers passing through the medial longitudinal
fasciculus to reach the thalamus that project in a diffuse way on the ventral posterolateral
nucleus.

Eferent connections from the brainstem over the vestibular organs arise from cells located
laterally to the external oculomotor nucleus.

Proprioceptive pathways
The position of the limbs and body in space depends basically on the information coming from
the neuromuscular spindle of the muscles, the Golgi tendinous organs in the tendons and the
Ruffini terminals in the joints.

The neuromuscular spindles are an important differentiation of muscle fibers. Most muscle
fibers of a muscle contract when they receive nerve impulses, these regular fibers are named
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extrafusal fibers. Between those fiubers there are some others that are enclosed in a capsule
loking as spindles that are not actually motor fibers but sensitive organs, these are the so-called
intrafusal fibers. These sensitive organs send to the CNS signals about the degree of tension in
the muscle and if the muscle has arrived to its goal. What allows to adapt the degree of tension
to the movement desired.

The spindles are bounded by an oval connective capsule that separates the intrafusal from the
extrafusals fibers. The intra fusal fibers are modified muscular cells that fuse into a sincitium
which concentrates the nuclei in the center of the fiber and becomes devoid of contractile actin
and myosin fibers, only the ends contain a few contractile fibers. There are two types of
intrafusal fibers the so-called nuclear bag and the nuclear chain. In the former, the central part
of the fiber has many nuclei and, in the second, the central part of the fiber contains nuclei
arranged in a row. Intrafusal fiber generate action potentials when the center (the one
containing the nuclei) is stretched. The way in which stretching occurs is important in the
generation of action potentials. The static fibers generate action potential asd a response to the
intensity of the stretching. In contrast, dynamic fibers respond to the speed of stretching.

The nuclear bag fibers can be either static or dynamic. In contrast, the nuclear chain fibers are
always static. A neuromuscular spindle typically contains two nuclear bag fibers (a static
dynamics) and approximately five nuclear chain fibers.

The neuromuscular spindle is innervated by two types of afferent axons: type IA or annulospiral
fibers anuloespirals and type II or tufted.

The IA fibers are large diameter myelinic fibers and fast conducting. Inside the spindle these
anulospiral fibers branch to arrive to nearly every every intrafusal fiber. Each terminal is coiled in
a spiral around the center of the fiber.

The secondary fibers (type II) fibers are also myelinic conducting fast but small diameter. The
secondary fibers innervate fibers in static nuclear bag and nuclear chain fibers, but not in
nuclear bag dynamic fibers.

On the other hand, neuromuscular spindles also receive efferent motor fibers (which come from
the ventral horn of the spinal cord. The motor neurons that control intrafusals fibers called
motor neurons and their axon fibers which are different from the regular αmotor neurons
which control extrafusal fibers. fibers innervate the distal part of the intrafusal fibers that keep
them tense. The tension of the intrafusal fiber depolarize it and activates the afferent fibers. At
this point there are two types of efferent fibers:

 Static fiber which innervates intrafusals nuclear chain fibers and static nuclear bag.

 Dynamic fibers which innervates nuclear bag dynamic fibers.

The system of the neuromuscular spindles, motor neurons and Golgi tendinous organs adjusts
the muscle tension to the desired movement to hold an object. So when trying to get something,
both α and  motor neurons are activated at the same time. The tension begins and activation of
intrafusal fibers forms a positive feedback circuit that activates the two motor neurons that keep
the tension until activation of the Golgi tendinous organ which triggers a negative feedback loop
which inhibits both motor neurons keeping the degree of tension.

Information about the degree of tension of the neuromuscular spindles, Golgi tendinous organs
as well as Ruffini’s terminals are being analyzed through the spinocerebellar pathways that
arrive to the spinocerebellum.
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In addition to be described in the topic about the cerebellum, a brief description of the pathway
must be done at this point. Prorpioceptive organs send information tom the spinal cord that
reach three main structures: the dorsal column the dorsal nucleus of Clarke and the
commissural nucleus. The dorsal column ascend through the spinal cord auntil arriving the
cuneate nucleus and then enter into the spinocerebellum through the juxtarestiformis organ.
The dorsal nucleus and the commissural organs send two kinds of projections. The direct dorsal
spinocerebellar pathway runs in the lateral cord without crossing until arriving to the pons and
entering in the inferior cerebellar peduncle into the spinocerebellum. The crossed anterior
spinocerebellar pathway cross the midline to arrive the anterior cord and ascend until the level
of the midbrain entering into the superior cerebellar peduncle and once inside the cerebellum
cross again to get into the ipsilateral cerebellum.

The spinal area of the cerebellum is the vermis and the paravermis and the corresponding
nucleus is the interpossitus. The cerebellum is in charge of the learning manual skills which
adjust the movement to an accurate goal.

Proprioception and vestibular representation in the cortex

Tracing methods have detailed the occurrence of vestibular projections from the rostral aspects
of the vestibular nuclei to the ventrobasal complex of the thalamus, namely the ventral
posterolateral, ventral posteromedial and ventral posteroinferior. This thalamic area also
receives information about proprioception together with the medial lemniscal afferents and
projects to the somatosensory cortex. Thus, in the somatosensory cortex there is overlapped
tactile, proprioceptive and vestibular information. On the other hand, the vestibular nuclei
receive corticval afferents arising from the prefrontal supplementary motor area (6a) which is
involved in oculomotor control and from areas 3aV which also receive optokinetic information.
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Diagram representing the relative position of the components of of the inner and middle ear on
the roof of the petrous portion of the temporal bone. a) ear, b) additus ad antrum mastoideum c)
antrum mastoideum d) hammer, e) anvil, f) chorda tympani nerve, g) cochlea, h) vestibulum, i)
superior semicircular canal, j) lateral semicircular canal, k) posterior semicircular canal, l)
ampulla of the semicircular duct, m) estatoacústic nerve, cranial nerve VIII
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Diagram representing the elements that makes up the membranous labyrinth and its
sensitive parts. a) superior vestibular ganglion, b) inferior vestibular ganglion ( together the
superior and inferior vestibular ganglia makes up the ganglion of Scarpa), c) spiral ganglion
and auditory nerve, d) cochlear duct, e) saculum with its sensitive side, f) utricle with its
sensitive part, g) ampulla of the superior semicircular duct, h) ampulla of the lateral
semicircular canal, i) ampulla of the posterior semicircular canal
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Diagram representing the location of the vestibular and cochlear nuclei in a dorsal view of the
brain stem (A), a transverse section at the pons-medulla transition (B) and a transverse section
at the same level in the rat stained for acetyl choline esterase (C). a) superior cerebellar
peduncle, b) middle cerebellar peduncle, c) inferior cerebellar peduncle, d) anterior ventral
cochlear nucleus, e) posterior ventral cochlear nucleus, f) dorsal cochlear nucleus, g) superior
vestibular nucleus, h) medial vestibular nucleus, i) lateral vestibular nucleus, j) inferior vestibular
nucleus, k) spinal trigeminal tract, l) nucleus of the trapezoid body, m), medial nucleus of the
superior olivary, n) lateral nucleus of the superior olivary complex, o) olivo-cochlear tract, p)
cochlear fibers, q) vestibular fibers, r) principal trigeminal nucleus, s), spinal trigeminal nucleus,
t) reticular formation.
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Distribution of vestibular fibers in the vestibular nuclei. Static stimuli coming from the utricle and
saculum are mainly represented in caudal aspects of the nuclei while dynamic projections
coming from senmicircular ducts targets the rostral aspects of the nuclei.
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Diagram representing the flow of information of the vestibulum-cerebellum. a) neuron of the

vestibular ganglion, b) medial vestibular nucleus, c) superior vestibular nucleus, d) lateral
vestibular nucleus, e) inferior vestibular nucleus, f) nodulus, g) flocsulus, h) fastigium nucleus, i)
nucleus interpossitus, j) dentate nucleus, k) unicinate tract (Russell), l) cerebello-reticular tracts,
m) medial vestibule-spinal tract, n) lateral vestibule-spinal tract, o) reticular formation
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Frontal eye field Visual associative cortex

Diagram representing the complex circuitry that govern movements of the eyes and head that
transmit the cranial nerves III, IV and VI coordinated with the movements of the neck of the XI
cranial nerve. The activity of oculomotor the accessory nuclei are coordinated by a number of
nuclei located around the medial longitudinal fasciculus. a) pretectum, b) superficial layer of the
superior colliculus c) deep layers of the superior colliculus, d) interstitial nucleus of Cajal and
Darckschwisch nucleus, e) inferior colliculus, f) accessory nucle nucleus of the common ocular
motor g) rostral part of the paramedian pontine reticular formation, h) common ocular motor
nucleus, y) trochlear nucleus, j) abducens nucleus, k) cerebellar floccular control of the
horitzontal gaze, m) caudal part of the paramedian pontine reticular formation, n) accessory
nucleus, o) medial longitudinal fasciculus.
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Diagram representing the muscular proprioceptive elements. a) neuromuscular spindle, b) α

motor fibers, c) tendinous Golgi organ, d) sensitive area, e) motor fiber γ, f) extrafusal fiber, g)
intrafusal fiber, m) conjunctive capsule, i) intrafusal fiber of nuclear chain, j) intrafusal fiber in
nuclear bag static, k) intrafusal fiber in nuclear bag dynamic, l) anulospiral fiber IA, m)
secondary tufted fiber II
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Diagram representing the flow of proprioceptive information in the spino-cerebellum. a)

propioreceptors of the neuromuscular spindles and Golgi tendinous organs, b) direct dorsal
spinocerebellar tract, c) dorsal columns, d) anterior crossed spinocerebellar tract, e) accessory
cuneatus nucleus, f) juxtarestiform tract, g) nucleus interpossitus, h) dentate nucleus, i) nucleus
fastigium, j) superior cerebellar peduncle, k) superior cerebellar peduncle, l) ventral lateral
nucleus of the thalamus, m) red nucleus, n) reticular formation, o) vestibular nuclei, p) vermis
and paravermis.