In this article we compare methods for temporal sequence learning (TSL) across the disciplines machine-control, classical conditioning, neuronal models for TSL as well as spiketiming dependent plasticity. This review will briefly introduce the most influential models and focus on two questions: 1) To what degree are reward-based (e.g. TD-learning) and correlation based (hebbian) learning related? and 2) How do the different models correspond to possibly underlying biological mechanisms of synaptic plasticity? We will first compare the different models in an open-loop condition, where behavioral feedback does not alter the learning. Here we observe, that reward-based and correlation based learning are indeed very similar. Machine-control is then used to introduce the problem of closed-loop control (e.g. “actor-critic architectures”). Here the problem of evaluative (“rewards”) versus nonevaluative (“correlations”) feedback from the environment will be discussed showing that both learning approaches are fundamentally different in the closed-loop condition. In trying to answer the second question we will compare neuronal versions of the different learning architectures to the anatomy of the involved brain structures (basal-ganglia, thalamus and

of reward expectation and reward expectation error in the putamen and caudate nucleus during stimulus-action-reward association learning. J Neurophysiol 95: 948–959, 2006. First published October 5, 2005; doi:10.1152/jn.00382.2005. To select appropriate behaviors leading to rewards, the brain needs to learn associations among sensory stimuli, selected behaviors, and rewards. Recent imaging and neural-recording studies have revealed that the dorsal striatum plays an important role in learning such stimulus-action-reward associations. However, the putamen and caudate nucleus are embedded in distinct cortico-striatal loop circuits, predominantly connected to motor-related cerebral cortical areas and frontal association areas, respectively. This difference in their cortical connections suggests that the putamen and caudate nucleus are engaged in different functional aspects of stimulus-action-reward association learning. To determine whether this is the case, we conducted an event-related and computational

Confronted with a rich sensory environment, the brain must learn statistical regularities across sensory domains to construct causal models of the world. Here, we used functional magnetic resonance imaging and dynamic causal modeling (DCM) to furnish neurophysiological evidence that statistical associations are learnt, even when task-irrelevant. Subjects performed an audio-visual target-detection task while being exposed to distractor stimuli. Unknown to them, auditory distractors predicted the presence or absence of subsequent visual distractors. We modeled incidental learning of these associations using a Rescorla--Wagner (RW) model. Activity in primary visual cortex and putamen reflected learning-dependent surprise: these areas responded progressively more to unpredicted, and progressively less to predicted visual stimuli. Critically, this prediction-error response was observed even when the absence of a visual stimulus was surprising. We investigated the underlying mechanism by embedding the RW model into a DCM to show that auditory to visual connectivity changed significantly over time as a function of prediction error. Thus, consistent with predictive coding models of perception, associative learning is mediated by prediction-error dependent changes in connectivity. These results posit a dual role for prediction-error in encoding surprise and driving associative plasticity.

In numerous and high-profile studies, researchers have recently begun to integrate computational models

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that neural correlates of the stimulus-action-reward association reside in the putamen, whereas a correlation with reward-prediction error was found largely in the caudate nucleus and ventral striatum. These non-uniform spatiotemporal distributions of neural correlates within the dorsal striatum were maintained consistently at various levels of task difficulty, suggesting a functional difference in the dorsal striatum between the putamen and caudate nucleus during stimulus-action-reward association learning.

A review of the current dopamine theories of schizophrenia reveals a likely imbalance between cortical and subcortical microcircuits due to an insufficient inhibitory brake, leading to a disruption of the dopamine system and the classic positive psychotic symptoms, negative symptoms and cognitive deficits associated with the disorder. Recent computational models have modelled the role of dopamine as a reward prediction error, using Temporal Difference and have successfully shown how these symptoms could arise from a disturbance to the dopamine system. We review these models in the light of dopamine theories of schizophrenia and highlight some of the major points that should be addressed by future computational models.

The desire to seek new and unfamiliar experiences is a fundamental behavioral tendency in humans and other species. In economic decision making, novelty seeking is often rational, insofar as uncertain options may prove valuable and advantageous in the long run. Here, we show that, even when the degree of perceptual familiarity of an option is unrelated to choice outcome, novelty nevertheless drives choice behavior. Using functional magnetic resonance imaging (fMRI), we show that this behavior is specifically associated with striatal activity, in a manner consistent with computational accounts of decision making under uncertainty. Furthermore, this activity predicts interindividual differences in susceptibility to novelty. These data indicate that the brain uses perceptual novelty to approximate choice uncertainty in decision making, which in certain contexts gives rise to a newly identified and quantifiable source of human irrationality.

Recent advances in the cognitive neuroscience of motivation and learning have demonstrated a critical role for midbrain dopamine and its targets in reward prediction. Converging evidence suggests that midbrain dopamine neurons signal a reward prediction error, allowing an organism to predict, and to act to increase, the probability of reward in the future. This view has been highly successful in accounting for a wide range of reinforcement learning phenomena in animals and humans. However, while current theories of midbrain dopamine provide a good account of behavior known as habitual or stimulus-response learning, we review evidence suggesting that other neural and cognitive processes are involved in motivated, goal-directed behavior. We discuss how this distinction resembles the classic distinction in the cognitive neuroscience of memory between nondeclarative and declarative memory systems, and discuss common themes between mnemonic and motivational functions. Finally, we present data demonstrating links between mnemonic processes and reinforcement learning. The past decade has seen a growth of interest in the cognitive neuroscience of motivation and reward. This is largely rooted in a series of neurophysiology studies of the response properties of dopamine-containing midbrain neurons in primates receiving reward (Schultz, 1998). The responses of these neurons were subsequently interpreted in terms of reinforcement learning, a computational framework for trial and error learning from reward (Houk, Adams, & Barto, 1995; Montague, Dayan, & Sejnowski, 1996; Schultz, Dayan, & Montague, 1997). Together with Both authors contributed equally to this article. We are most grateful to Shanti Shanker for assistance with data collection, to Anthony Wagner for generously allowing us to conduct the experiment reported here in his laboratory, and to Alison Adcock, Lila Davachi, Peter Dayan, Mark

We used behavioral and functional magnetic resonance imaging (fMRI) methods to probe the cerebral organization of a simple logical deduction process. Subjects were engaged in a motor trialand-error learning task, in which they had to infer the identity of an unknown 4-key code. The design of the task allowed subjects to base their inferences not only on the feedback they received but also on the internal deductions that it afforded (autoevaluation). fMRI analysis revealed a large bilateral parietal, prefrontal, cingulate, and striatal network that activated suddenly during search periods and collapsed during ensuing periods of sequence repetition. Fine-grained analyses of the temporal dynamics of this search network indicated that it operates according to near-optimal rules that include 1) computation of the difference between expected and obtained rewards and 2) anticipatory deductions that predate the actual reception of positive reward. In summary, the dynamics of effortful mental deduction can be tracked with fMRI and relate to a distributed network engaging prefrontal cortex and its interconnected cortical and subcortical regions.