Pitching the head while rotating (PWR) combines periodic activation
of the semicircular canals and the otoliths to generate pitch
and roll eye deviations and continuous horizontal nystagmus.
Monkeys were tested after individual pairs of semicircular canals
were plugged and single units were recorded in the vestibular
nerve while the animals were sinusoidally pitched 20-40 deg
about a spatial horizontal axis with 5- and 16-s periods and
simultaneously rotated about a spatial vertical axis at 30-120
deg/s. As previously shown, the steady-state horizontal response
disappeared after plugging the vertical semicircular canals,
but was maintained when the lateral canals were plugged. When
the left anterior and right posterior canal (LARP) pair was
left intact, the steady-state response depended on the axis
about which the pitching took place. When the axis was normal
to the LARP plane, there was no steady-state response. When
the pitching axis was perpendicular to the LARP normal, the
response was maximal. Firing rates of otolith units were approximately
in phase with pitch position, and the addition of rotation about
a vertical axis did not change the response. Lateral canal units
did not have a steady-state modulation during pitch or constant
velocity rotation. During PWR, they oscillated at twice the
pitch frequency. This corresponded to the frequency at which
the canal was maximally activated as it aligned with the plane
of rotation. The amplitude of modulation increased proportionally
to rotational velocity, but the phase remained the same. These
characteristics were unchanged during roll while rotating (RWR),
which induces little continuous nystagmus. Anterior and posterior
canal units were maximally excited near pitch-velocity maxima
and minima, respectively, during pure pitching. During PWR,
however, the phases of both components simultaneously shifted
toward each other and toward being in phase with otolith units.
The peak excitation tended toward a forward-pitch position when
the rotation was to the ipsilateral side, and toward a backward
pitch position when the rotation was to the contralateral side.
With 120-deg/s rotation during a 16-s pitch period, the phase
difference between anterior and posterior canal units was as
small as 17 deg. These data support the postulate that the correlation
between vertical canal and otolith units is the critical factor
in generating continuous unidirectional horizontal nystagmus
during PWR.