In some experiments, targets moved in the four cardinal directions with speeds of 5, 10, or 20°/s in different trials. Each learning experiment consisted of a baseline block and a learning block. During

the baseline block (80 to 100 trials), the target moved at 20°/s in one of selleck two opposing cardinal directions, designated the probe (55% of the trials) and control directions (45%). In the learning block, (250 to 300 trials) the pursuit target also initially moved in either the probe (55%) or control (45%) directions; however, targets moving in the probe direction had an 82% chance of adopting a 30°/s orthogonal velocity component at a fixed time after the onset of target motion. The direction and timing of the instructive stimulus was fixed for a given learning block. In some recording sessions, we performed an additional learning experiment after residual Selleck Tariquidar behavioral learning had been extinguished with a second baseline block (100–150 trials) or a two-block sequence of learning in the opposite direction (50-100 trials) followed by a baseline block (50 trials). The residual eye velocity measured after

the two reversal procedures averaged 27.7% (SD: 30.8%, range: 61.7% to −34.1%) of the original learned response after a baseline block and −1.3% (SD: 16.3%, range: 47.9% to −33.1%) after a learning block in the opposite direction and another baseline block. For 21 neurons, we followed the reversal procedure with a mimic experiment, which consisted of a baseline block followed by a mimic block. The mimic block featured mimic trials designed to evoke an eye velocity with the same time course and trajectory as the learned component of eye movement but without any learning. To prevent learning during the mimic block, we counterbalanced mimic trials in the learning direction with trials that contained the same target perturbation in the opposite direction.

Trials were examined individually by eye to identify the onset and offset times of any saccades; we replaced the intervening eye velocity with a linear interpolation whose the endpoints were the eye velocity values at the onset and offset of the saccade. We quantified the magnitude of neural learning in the interval from 100 ms after the onset of target motion to 70 ms after the instruction time, as the difference in mean spike count between the learning trials in the learning block and the probe trials from the baseline block. Neural responses are reported as firing rates, obtained by dividing the spike counts by the duration of the analysis intervals. We verified that all analyses produced similar results if the firing rate changes were converted to Z-scores.