We would rather decline an effortful option, but when compelled, will move only slowly to harvest. Why should economic variables such as reward and effort affect movement vigor? In theory, both our decisions and our movements contribute to a measure of fitness in which the objective is to maximize rewards minus efforts, divided by time. To explore this idea, we engaged marmosets in a foraging task in which on each trial they decided whether to work by making saccades to visual targets, thus accumulating food, or to harvest by licking what they had earned. We varied the effort cost of harvest by moving the food tube with respect to the mouth. Theory predicted that the subjects should respond to the increased effort costs by working longer, stockpiling food before commencing harvest, but reduce their movement vigor to conserve energy. Indeed, in response to the increased effort costs of harvest, marmosets increased their work duration but reduced their movement vigor. These changes in decisions and movements coincided with changes in pupil size. As the effort cost of harvest declined, work duration decreased, the pupils dilated, and lick and saccade vigor increased. Thus, when acquisition of reward became effortful, there was a global change in the state of the brain: the pupils constricted, the decisions exhibited delayed gratification, and the movements displayed reduced vigor.
Why do economic variables such as reward and effort affect both the decision-making and the motor-control circuits of the brain? Our results suggest that as the brainstem neuromodulatory circuits that control pupil size respond to effort costs, they alter computations in the brain regions that control decisions, encouraging work and delaying gratification, and the brain regions that control movements, suppressing energy expenditure and reducing vigor. This coordinated response may improve a variable relevant to fitness: the capture rate.