Interaction experiments on the responses evoked in Purkinje cells by climbing fibres.

1. The uniquely powerful excitatory synaptic action of a single climbing fibre on a Purkinje cell in the cerebellum of the cat was tested during the intense and prolonged inhibitory action produced by the parallel fibre, basket and stellate cell system. There was depression of the later spike discharges, but the initial discharge was never suppressed.2. With intracellular recording the excitatory post-synaptic potential was depressed during the initial phase (about 10 msec) of the inhibitory action, but there was a later increase with a time course resembling the latter part of the inhibitory hyperpolarization. An explanation of these and other effects is given in terms of conventional ideas of excitatory and inhibitory synaptic interaction.3. These observations on single Purkinje cells, particularly with intracellular recording, have helped in formulating a provisional explanation of the finding that during inhibition there is an increase in the negative field potential evoked by a climbing fibre volley.4. The excitatory action of a climbing fibre synapse is shown to be greatly depressed immediately after a preceding activation and recovery takes hundreds of milliseconds. By the collision technique it is shown that the same climbing fibre is activated by inferior olive and juxta-fastigial stimulation.5. With rapid repetitive activation there was initially a progressive decline in the effectiveness of each successive impulse, but a steady level was soon reached. On cessation of a tetanus of twenty or more impulses there was a delayed recovery of the depolarization, which suggests a continued action of the accumulated transmitter.6. With extracellular recording repetitive spike initiation continued with stimulation frequencies as high as 100/sec, but at still higher frequencies spikes were depressed by the intense synaptically evoked depolarization. On cessation of the stimulation after-discharge often developed as the depolarization declined. The prolonged after-discharges following severe tetani suggest that there is a very effective accumulation of the transmitter.