Previous animal experiments have shown that serotonin is involved in the control of impulsive choice, as characterized by high preference for small immediate rewards over larger delayed rewards. Previous human studies under serotonin manipulation, however, have been either inconclusive on the effect on impulsivity or have shown an effect in the speed of action-reward learning or the optimality of action choice. Here, we manipulated central serotonergic levels of healthy volunteers by dietary tryptophan depletion and loading. Subjects performed a "dynamic" delayed reward choice task that required a continuous update of the reward value estimates to maximize total gain. By using a computational model of delayed reward choice learning, we estimated the parameters governing the subjects' reward choices in low-, normal, and high-serotonin conditions. We found an increase of proportion in small reward choices, together with an increase in the rate of discounting of delayed rewards in the low-serotonin condition compared with the control and high-serotonin conditions. There were no significant differences between conditions in the speed of learning of the estimated delayed reward values or in the variability of reward choice. Therefore, in line with previous animal experiments, our results show that low-serotonin levels steepen delayed reward discounting in humans. The combined results of our previous and current studies suggest that serotonin may adjust the rate of delayed reward discounting via the modulation of specific loops in parallel corticobasal ganglia circuits.
BackgroundThe ability to select an action by considering both delays and amount of reward outcome is critical for maximizing long-term benefits. Although previous animal experiments on impulsivity have suggested a role of serotonin in behaviors requiring prediction of delayed rewards, the underlying neural mechanism is unclear.Methodology/Principal FindingsTo elucidate the role of serotonin in the evaluation of delayed rewards, we performed a functional brain imaging experiment in which subjects chose small-immediate or large-delayed liquid rewards under dietary regulation of tryptophan, a precursor of serotonin. A model-based analysis revealed that the activity of the ventral part of the striatum was correlated with reward prediction at shorter time scales, and this correlated activity was stronger at low serotonin levels. By contrast, the activity of the dorsal part of the striatum was correlated with reward prediction at longer time scales, and this correlated activity was stronger at high serotonin levels.Conclusions/SignificanceOur results suggest that serotonin controls the time scale of reward prediction by differentially regulating activities within the striatum.
Critical to our many daily choices between larger delayed rewards, and smaller more immediate rewards, are the shape and the steepness of the function that discounts rewards with time. Although research in artificial intelligence favors exponential discounting in uncertain environments, studies with humans and animals have consistently shown hyperbolic discounting. We investigated how humans perform in a reward decision task with temporal constraints, in which each choice affects the time remaining for later trials, and in which the delays vary at each trial. We demonstrated that most of our subjects adopted exponential discounting in this experiment. Further, we confirmed analytically that exponential discounting, with a decay rate comparable to that used by our subjects, maximized the total reward gain in our task. Our results suggest that the particular shape and steepness of temporal discounting is determined by the task that the subject is facing, and question the notion of hyperbolic reward discounting as a universal principle.
We investigated the event-related power decrease (event-related desynchronization: ERD) of the alpha bands associated with the anticipation of affective images. Participants (n=19) were presented with emotionally positive or negative images under different anticipatory conditions, and their brain responses were recorded using magnetoencephalography (MEG). In the Affective Cue conditions, the cue stimulus indicated the emotional valence (positive or negative) of the image. In the Null Cue condition, the cue stimulus did not include any information about the valence of the image, and in the No Cue condition, the affective image was presented without a preceding cue. The cues in the affective and null conditions were followed by emotional images. During the anticipation period for the affective image, the alpha ERD preceding an anticipated negative image was larger than that preceding an anticipated positive image; this effect had an occipital dominance. Furthermore, during the anticipation period, the lower-2-alpha ERD of the right frontal area showed the same result. These results demonstrate that anticipation of negative stimuli induced alpha ERD in both the visual and the right frontal cortex, indicating that top-down modulation may be provided by the right frontal cortex to the visual cortex.
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