Starting from macroscopic and microscopic facts of cerebellar
histology, we propose a new functional interpretation that may
elucidate the role of the cerebellum in movement control. The idea
is that the cerebellum is a large collection of individual lines
(Eccles's “beams”: Eccles et al. 1967a) that
respond specifically to certain sequences of events in the input
and in turn produce sequences of signals in the output. We believe
that the sequence-in/sequence-out mode of operation is as typical
for the cerebellar cortex as the transformation of sets into sets of
active neurons is typical for the cerebral cortex, and that both
the histological differences between the two and their reciprocal
functional interactions become understandable in the light of this
dichotomy. The response of Purkinje cells to sequences of stimuli
in the mossy fiber system was shown experimentally by Heck on
surviving slices of rat and guinea pig cerebellum. Sequential
activation of a row of eleven stimulating electrodes in the granular
layer, imitating a “movement” of the stimuli along
the folium, produces a powerful volley in the parallel fibers that
strongly excites Purkinje cells, as evidenced by intracellular
recording. The volley, or “tidal wave,” has maximal
amplitude when the stimulus moves toward the recording site at
the speed of conduction in parallel fibers, and much smaller
amplitudes for lower or higher “velocities.” The
succession of stimuli has no effect when they “move”
in the opposite direction. Synchronous activation of the stimulus
electrodes also had hardly any effect. We believe that the sequences
of mossy fiber activation that normally produce this effect in the
intact cerebellum are a combination of motor planning relayed to
the cerebellum by the cerebral cortex, and information about ongoing
movement, reaching the cerebellum from the spinal cord. The output
elicited by the specific sequence to which a “beam” is
tuned may well be a succession of well timed inhibitory volleys
“sculpting” the motor sequences so as to adapt them
to the complicated requirements of the physics of a multijointed
system.