As already mentioned, the orientation of the hair cell bundles is organized relative to

the striola, which demarcates the overlying layer of otoconia (see Figure 14.4A). The striola
forms an axis of mirror symmetry such that hair cells on opposite sides of the striola have
opposing morphological polarizations. Thus, a tilt along the axis of the striola will excite the
hair cells on one side while inhibiting the hair cells on the other side. The saccular macula is
oriented vertically and the utricular macula horizontally, with a continuous variation in the
morphological polarization of the hair cells located in each macula (as shown in Figure
14.4C, where the arrows indicate the direction of movement that produces excitation).
Inspection of the excitatory orientations in the maculae indicates that the utricle responds to
movements of the head in the horizontal plane, such as sideways head tilts and rapid lateral
displacements, whereas the sacculus responds to movements in the vertical plane (up-down
and forward-backward movements in the sagittal plane). Note that the saccular and utricular
maculae on one side of the head are mirror images of those on the other side. Thus, a tilt of
the head to one side has opposite effects on corresponding hair cells of the two utricular
maculae. This concept is important in understanding how the central connections of the
vestibular periphery mediate the interaction of inputs from the two sides of the head (see next
section).

Extensor activity is induced on the side to which the head is inclined, and
flexor activity is induced on the opposite side. The head movement opposes
the movement registered by the vestibular system.

The output neurons of the VSR are the anterior horn cells of the spinal cord
gray matter, which drive skeletal muscle. However, the connection between
the vestibular nuclear complex and the motor neurons is more complicated
than for the VOR.

The VSR has a much more difficult task than the VOR, because there are
multiple strategies that can be used to prevent falls, which involve entirely
different motor synergies. For example, when shoved from behind, one’s
centre of gravity might become displaced anteriorly. In order to restore
Vestibular Spinal Reflex (VSR)
The VSR stabilises the body. As an example of a vestibulospinal reflex, let
us examine the sequence of events involved in generating a labyrinthine
reflex.

1. When the head is tilted to one side, both the canals and otoliths are
stimulated. Endolymphatic flow deflects the cupula and shear force
deflects hair cells within the otoliths.
2. The vestibular nerve and vestibular nucleus are activated.
3. Impulses are transmitted via the lateral and medial vestibulospinal
tracts to the spinal cord.

“balance,” one might (1) plantarflex at the ankles; (2) take a step; (3) grab
for support; or (4) use some combination of all three activities.

The VSR also has to adjust limb motion appropriately for the position of
the head on the body. The VSR must also use otolith input, reflecting linear
motion, to a greater extent than the VOR.  The eyes can only rotate and
thus can do little to compensate for linear motion, whilst the body can both
rotate AND translate.