Macular input to the cerebellar nodulus

The ability to orient and navigate through the terrestrial environment represents a computational challenge common to all vertebrates. It arises because motion sensors in the inner ear, the otolith organs, and the semicircular canals transduce self-motion in an egocentric reference frame. As a result, vestibular afferent information reaching the brain is inappropriate for coding our own motion and orientation relative to the outside world. Here we show that cerebellar cortical neuron activity in vermal lobules 9 and 10 reflects the critical computations of transforming head-centered vestibular afferent information into earth-referenced self-motion and spatial orientation signals. Unlike vestibular and deep cerebellar nuclei neurons, where a mixture of responses was observed, Purkinje cells represent a homogeneous population that encodes inertial motion. They carry the earth-horizontal component of a spatially transformed and temporally integrated rotation signal from the semicircular canals, which is critical for computing head attitude, thus isolating inertial linear accelerations during navigation.

In a first series of experiments, recordings were obtained from cat abducens and trochlear motorneurons and from axons of secondary vestibular neurons terminating in these motor nuclei, and the effects of cerebellar nodulus stimulation on utricular- and canal-evoked responses in these neurons were studied.

Ultricular activation of vestibular axons recorded in the ipsilateral VIth and contralateral IVth nuclei was probably monosynaptically inhibited by nodular stimulation provided conditioning-test intervals were in the range between 0–10 ms and the test stimuli were close to threshold intensities. Of the vestibular axons activated by stimulation of the semicircular canal nerves only those evoked by the horizontal canal stimulation and recorded in the ipsilateral VIth nucleus were weakly inhibited. When the vestibular stimuli were strong enough to produce clear field potentials in the motor nuclei and/or postsynaptic potentials in motorneurons, nodular stimulation had practically no effect on their amplitudes. It is concluded that inhibition of vestibuloocular transmission is weak as compared to floccular inhibition studied previously.

In a second series of experiments, recordings were obtained from vestibular neurons which were activated antidromically and/or transsynaptically by stimulation of the contralateral fastigial nucleus, and the effects of ipsilateral nodular stimulation on these responses were studied. It was found that nodular stimulation inhibited both antidromic as well as transsynaptic fastigial activations of vestibular neurons. Most of these vestibular neurons were located in the descending vestibular nucleus and received polysynaptic vestibular and spinal inputs. It is concluded that in addition to its weak inhibitory effect on vestibuloocular transmission the nodulus exerts a powerful inhibition on vestibular neurons transmitting vestibular and spinal inputs to cerebellar nuclei and/or cortex.

It is suggested that the nodulus controls cerebellar projecting vestibular neurons which carry vestibular and spinal information to the cerebellum. The vestibular, proprioceptive and visual information which is present in the nodulus may aid the role of the nodulus in controlling body posture.