SENSORY PROCESSING AND COROLLARY DISCHARGE EFFECTS IN THE MORMYROMAST REGIONS OF THE MORMYRID ELECTROSENSORY LOBE .1. FIELD POTENTIALS, CELLULAR-ACTIVITY IN ASSOCIATED STRUCTURES

1. This is the first of a series of papers on the electrosensory lobe and closely associated structures in electric fish of the family Mormyridae. The study describes the neuronal responses to sensory stimuli and to corollary discharge signals associated with the motor command that drives the electric organ discharge (EOD). The study is focused on the regions of the electrosensory lobe where primary afferent fibers from mormyromast electroreceptors terminate. 2. This first paper of the series describes the field potentials in the caudal lobe of the cerebellum and in the electrosensory lobe. It also describes the different types of unit activity in the caudal lobe of the cerebellum. Granule cells of the caudal lobe of the cerebellum provide the parallel fibers for most of the molecular layer of the electrosensory lobe. Determination of the input and responses of these cells is therefore an important part of the effort to understand the electrosensory lobe. 3. Corollary discharge field potentials evoked by the EOD motor command are very prominent in the caudal lobe of the cerebellum and in the electrosensory lobe. The potentials indicate that corollary discharge excitation affects first the granule cells of the caudal lobe and then, a few milliseconds later, the deeper cellular layers of the electrosensory lobe. The prominence and complexity of the field potentials indicate that corollary discharge signals have an important and varied role in the processing of electrosensory information by the mormyrid electrosensory lobe. 4. The field potentials evoked by electrosensory stimuli suggest that direct primary afferent excitation is limited to the granule and intermediate layers of the electrosensory lobe, as is indicated also by anatomic studies. 5. Proprioceptive units are the most common type of unit recorded in the granule cell region of the caudal lobe of the cerebellum (eminentia granularis posterior). These units have a regular discharge rate that changes tonically in response to slight bending of the trunk, bending of the tail, or bending of individual fins. Proprioceptive input will have a strong effect on the molecular layer of the electrosensory lobe and will thus modulate the responses of electrosensory lobe cells to electrosensory stimuli. Such proprioceptive input to the electrosensory lobe would allow the expected effects of body position changes to be accounted for in the processing of electrosensory information. 6. Units with stereotyped, short-latency corollary discharge bursts to the EOD motor command were the next most common type of unit in the eminentia granularis posterior. These corollary discharge units were not affected by sensory stimuli. These units probably mediate the short-latency corollary discharge excitation of the granule cells that was also indicated by the field-potential recordings. 7. Additional types of units in the eminentia granularis posterior included the following: corollary discharge units with long latencies and variable bursts to the EOD motor command; corollary discharge units with stereotyped pauses of various durations following the EOD motor command; and mechanical lateral line units responding to water movement along the trunk. These types of units will also affect the electrosensory lobe via the axons of granule cells that end as parallel fibers in the molecular layer. 8. Purkinje cells of the caudal lobe molecular layer could be identified by their characteristic climbing fiber responses. Both the climbing fiber responses and the ordinary spikes of Purkinje cells were affected by corollary discharge signals and by electrosensory stimuli. 9. The physiological results and the close anatomic parallels between the caudal lobe of the cerebellum and the electrosensory lobe indicate that both the granule cell region and the molecular layer of the caudal lobe have roles in the processing of electrosensory information.