Distinct error-correcting and incidental learning of location relative to landmarks and boundaries

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Abstract
Associative reinforcement provides a powerful explanation of learned behavior. However, an unproven but long-held conjecture holds that spatial learning can occur incidentally rather than by reinforcement. Using a carefully controlled virtual-reality object-location memory task, we formally demonstrate that locations are concurrently learned relative to both local landmarks and local boundaries but that landmark-learning obeys associative reinforcement (showing "overshadowing" and "blocking" or "learned irrelevance"), whereas boundary-learning is incidental, showing neither overshadowing nor blocking nor learned irrelevance. Crucially, both types of learning occur at similar rates and do not reflect differences in levels of performance, cue salience, or instructions. These distinct types of learning likely reflect the distinct neural systems implicated in processing of landmarks and boundaries: the striatum and hippocampus, respectively [Doeller CF, King JA, Burgess N (2008) Proc Natl Acad Sci USA 105:5915-5920]. In turn, our results suggest the use of fundamentally different learning rules by these two systems, potentially explaining their differential roles in procedural and declarative memory more generally. Our results suggest a privileged role for surface geometry in determining spatial context and support the idea of a "geometric module," albeit for location rather than orientation. Finally, the demonstration that reinforcement learning applies selectively to formally equivalent aspects of task-performance supports broader consideration of two-system models in analyses of learning and decision making.