The insula, a cortical brain region that is known to encode information about autonomic, visceral, and olfactory functions, has recently been shown to encode information during reward-seeking tasks in both single neuronal recording and functional magnetic resonance imaging studies. To examine the reward-related activation, we recorded from 170 single neurons in anterior insula of 2 monkeys during a multitrial reward schedule task, where the monkeys had to complete a schedule of 1, 2, 3, or 4 trials to earn a reward. In one block of trials a visual cue indicated whether a reward would or would not be delivered in the current trial after the monkey successfully detected that a red spot turned green, and in other blocks the visual cue was random with respect to reward delivery. Over one-quarter of 131 responsive neurons were activated when the current trial would (certain or uncertain) be rewarded if performed correctly. These same neurons failed to respond in trials that were certain, as indicated by the cue, to be unrewarded. Another group of neurons responded when the reward was delivered, similar to results reported previously. The dynamics of population activity in anterior insula also showed strong signals related to knowing when a reward is coming. The most parsimonious explanation is that this activity codes for a type of expected outcome, where the expectation encompasses both certain and uncertain rewards.

The dorsal raphe nucleus is the major source of serotonin in the brain. It is connected to brain regions related to reward processing, and the neurons show activity related to predicted reward outcome. Clinical observations also suggest that it is important in maintaining alertness and its apparent role in addiction seems to be related to reward processing. Here, we examined whether the neurons in dorsal raphe carry signals about reward outcome and task progress during multitrial schedules. We recorded from 98 single neurons in dorsal raphe of two monkeys. The monkeys perform one, two, or three visual discrimination trials (schedule), obtaining one, two, or three drops of liquid. In the valid cue condition, the length and brightness of a visual cue indicated schedule progress and reward amount, respectively. In the random cue condition, the visual cue was randomly presented with respect to schedule length and reward amount. We found information encoded about (1) schedule onset, (2) reward expectation, (3) reward outcome, and (4) reward amount in the mean firing rates. Information theoretic analysis showed that the temporal variation of the neuronal responses contained additional information related to the progress of the schedule toward the reward rather than only discriminating schedule onset or reward/no reward. When considered in light of all that is known about the raphe in anatomy, physiology, and behavior, the rich encoding about both task progress and predicted reward outcome makes the raphe a strong candidate for providing signals throughout the brain to coordinate persistent goal-seeking behavior.

The recorded responses of single neurons often vary considerably in the numbers of spikes emitted across repeats of a single experimental condition. Because of this irregularity and for theoretical convenience the responses are often approximated using a Poisson process. However, it has been frequently pointed out that many details of the responses, including the distribution of spike counts across similar trials, are not consistent with a Poisson process, even an inhomogeneous one. Wiener and Richmond (2003, J Neurosci 23:2394-2406) showed that the spike count distributions could usually be fitted nicely by mixtures of a few (1-3) Poisson distributions, a step they regarded as a computational convenience. Now, we find that a substantial proportion (47%) of the neuronal responses from anterior cingulate cortex, which we conceptualize as part of a system related to the balance between work and reward, have responses with multimodal firing rate distributions. When these distributions are modeled as mixtures of Poisson distributions, the proportions of the different Poisson distributions are related to behavioral state, and might be related to cognitive factors. This suggests that the neurons undergo behaviorally-related mode changes.

Choice reflects the values of available alternatives; more valuable options are chosen more often than less valuable ones. Here we studied whether neuronal responses in orbitofrontal cortex (OFC) reflect the value difference between options, and whether there is a causal link between OFC neuronal activity and choice. Using a decision-making task where two visual stimuli were presented sequentially, each signifying a value, we showed that when the second stimulus appears many neurons encode the value difference between alternatives. Later when the choice occurs, that difference signal disappears and a signal indicating the chosen value emerges. Pharmacological inactivation of OFC neurons coding for choice-related values increases the monkey’s latency to make a choice and the likelihood that it will choose the less valuable alternative, when the value difference is small. Thus, OFC neurons code for value information that could be used to directly influence choice.

BackgroundThe value of a predicted reward can be estimated based on the conjunction of both the intrinsic reward value and the length of time to obtain it. The question we addressed is how the two aspects, reward size and proximity to reward, influence the responses of neurons in rostral anterior cingulate cortex (rACC), a brain region thought to play an important role in reward processing.Methods and FindingsWe recorded from single neurons while two monkeys performed a multi-trial reward schedule task. The monkeys performed 1–4 sequential color discrimination trials to obtain a reward of 1–3 liquid drops. There were two task conditions, a valid cue condition, where the number of trials and reward amount were associated with visual cues, and a random cue condition, where the cue was picked from the cue set at random. In the valid cue condition, the neuronal firing is strongly modulated by the predicted reward proximity during the trials. Information about the predicted reward amount is almost absent at those times. In substantial subpopulations, the neuronal responses decreased or increased gradually through schedule progress to the predicted outcome. These two gradually modulating signals could be used to calculate the effect of time on the perception of reward value. In the random cue condition, little information about the reward proximity or reward amount is encoded during the course of the trial before reward delivery, but when the reward is actually delivered the responses reflect both the reward proximity and reward amount.ConclusionsOur results suggest that the rACC neurons encode information about reward proximity and amount in a manner that is dependent on utility of reward information. The manner in which the information is represented could be used in the moment-to-moment calculation of the effect of time and amount on predicted outcome value.