Figure 3.

Trace conditioning with probabilistic rewards. (a) An illustration of one trial of the delay conditioning task of Fiorillo et al. [15]. A trial consists of a 2-second visual stimulus, the offset of which coincides with the delivery of the juice reward, if such a reward is programmed according to the probability associated with the visual cue. In unrewarded trials the stimulus terminated without a reward. In both cases an inter-trial interval of 9 seconds on average separates trials. (b) An illustration of one trial of the trace conditioning task of Morris et al. [16]. The crucial difference is that there is now a substantial temporal delay between the offset of the stimulus and the onset of the reward (the "trace" period), and no external stimulus indicates the expected time of reward. This confers additional uncertainty as precise timing of the predicted reward must be internally resolved, especially in unrewarded trials. In this task, as in [15], one of several visual stimuli (not shown) was presented in each trial, and each stimulus was associated with a probability of reward. Here, also, the monkey was requested to perform an instrumental response (pressing the key corresponding to the side in which the stimulus was presented), the failure of which terminated the trial without a reward. Trials were separated by variable inter-trial intervals. (c,d) DA firing rate (smoothed) relative to baseline, around the expected time of the reward, in rewarded trials (c) and in unrewarded trials (d). (c,d) Reprinted from [16] ©2004 with permission from Elsevier. The traces imply an overall positive response at the expected time of the reward, but with a very small, or no ramp preceding this. Similar results were obtained in a classical conditioning task briefly described in [15], which employed a trace conditioning procedure, confirming that the trace period, and not the instrumental nature of the task depicted in (b) was the crucial difference from (a).