FICTIVE SWIMMING ELICITED BY ELECTRICAL-STIMULATION OF THE MIDBRAIN IN GOLDFISH

1. We developed a fictive swimming preparation of goldfish that will allow us to study the cellular basis of interactions between swimming and escape networks in fish. 2. Stimulation of the midbrain in decerebrate goldfish produced rhythmic alternating movements of the body and tail similar to swimming movements. The amplitude and frequency of the movements were dependent on stimulus strength. Larger current strengths or higher frequencies of stimulation produced larger-amplitude and/or higher-frequency movements. Tail-beat frequency increased roughly linearly with current strength over a large range, with plateaus in frequency sometimes evident at the lowest and highest stimulus strengths. 3. Electromyographic (EMG) recordings from axial muscles on opposite sides at the same rostrocaudal position showed that stimulation of the midbrain led to alternating EMG bursts, with bursts first on one side, then the other. These bursts occurred at a frequency equal to the tail-beat frequency and well below the frequency of brain stimulation. EMG bursts recorded from rostral segments preceded those recorded from caudal segments on the same side of the body. The interval between individual spikes within EMG bursts sometimes corresponded to the interval between brain stimuli. Thus, whereas the frequency of tail beats and EMG bursts was always much slower than the frequency of brain stimulation, there was evidence of individual brain stimuli in the pattern of spikes within bursts. 4. After paralyzing fish that produced rhythmic movement on midbrain stimulation, we monitored the motor output during stimulation of the midbrain by using extracellular recordings from spinal motor nerves. We characterized the motor pattern in detail to determine whether it showed the features present in the motor output of swimming fish. The fictive preparations showed all of the major features of the swimming motor pattern recorded in EMGs from freely swimming fish. 5. The motor nerves, like the EMGs produced by stimulating midbrain, showed rhythmic bursting at a much lower frequency than the brain stimulus. Bursts on opposite sides of the body alternated. The frequency of bursting ranged from 1.5 to 13.6 Hz and was dependent on stimulus strength, with higher strengths producing faster bursting. Activity in rostral segments preceded activity in caudal ones on the same side of the body. Some spikes within bursts of activity occurred at the same frequency as the brain stimulus, but individual brain stimuli were not as evident as those seen in some of the EMGs. 6. The duration of bursts of activity in a nerve was positively and linearly correlated with the time between successive bursts (cycle time). Increases in cycle time were associated with increases in burst duration. Burst duration was usually a constant fraction of cycle time, with bursts occupying on average 50.6% of cycle time. 7. The rostrocaudal delay was linearly and positively correlate with cycle time. Larger cycle times were associated with large delays. The delay was usually a constant fraction of cycle time at different burst frequencies. It averaged 2.1% of a cycle per body segment. This implies that at any point in time there is roughly 60% of a wave of activity along the 29- to 30-segment body and tail of the fish. 8. We conclude that the motor pattern produced by midbrain stimulation in the paralyzed goldfish was a pattern that would produce swimming if the fish was free to move. The development and characterization of this fictive swimming preparation sets the stage for future studies of interactions between the spinal network for swimming and the well-studied network responsible for escape movements in goldfish.