Exhumed from thought: basal ganglia and response learning in the plus-maze

The plus-maze apparatus figured prominently in the historical debate between cognitive and stimulus-response habit learning theorists concerned with the fundamental question of "what" animals learn. An important feature of this task is that variants of the training procedure can be arranged to allow for an assessment of the relative use of cognitive/place or habit/response learning mechanisms. This brief review describes findings from several neurobiological studies published primarily over the past decade that have re-introduced the plus-maze to investigate the role of the dorsal striatum in learning and memory. Converging evidence from research using brain lesion, pharmacological, and molecular/genetic approaches is described supporting the hypothesis that the dorsolateral striatum plays a selective role in response learning in the plus-maze. Within a multiple systems framework of memory organization, factors that can influence the relative use of place and response learning in the plus-maze are also considered, including the nature of the visual environment, reinforcement/training parameters, and emotional state of the organism. Response learning in the plus-maze may be considered an exemplar task useful for investigating the neurobiological bases of dorsal striatal involvement in habit learning and memory. This mnemonic function of the dorsal striatum generalizes across several sensory modalities and mammalian species, including humans.