Our goal was to study potential substrates for cortical modulation of vesti
bular reflexes in the cat. In initial experiments, injections of wheat-germ
-agglutinate-horseradish-peroxidase into Deiters' nucleus and the rostral d
escending nucleus revealed bilateral colonies of retrogradely filled neuron
s in cortical areas 6, 2, and 3a (about 60 cells per colony). In cats anest
hetized with chloralose-urethane, we stimulated areas 2 and 3a with trains
of pulses while recording from ipsilateral vestibular-nucleus neurons, whic
h were characterized by their responses to sinusoidal tilts and tested for
the presence of antidromic responses to stimulation of the upper cervical c
ord. A majority of the neurons was affected by cortical stimulation, showin
g either facilitation, inhibition, or a mixture of the two. Stimulation in
area 2 was more effective than stimulation in area 3a. Despite the anatomic
presence of direct cortico-vestibular projections, properties of facilitat
ion and inhibition suggest that both were evoked by polysynaptic pathways.
Cortical effects were broadly distributed to vestibular neurons without reg
ard to responses of these neurons to sinusoidal tilts. There was no signifi
cant difference between effects on lateral and medial vestibulospinal tract
neurons, but, as a group, vestibulospinal neurons were much more likely to
be affected by cortical stimulation than neurons not antidromically activa
ted from the C2 segment. We conclude that, by their influence on vestibulos
pinal neurons, neurons in cortical areas 2 and 3a should be able to modulat
e, in behaving animals, vestibular reflexes acting on the neck and limbs.