Generalization via superposition: combined effects of mixed reference frame representations for explicit and implicit learning in a visuomotor adaptation task
Studies on generalization of learned visuomotor perturbations have generally focused on whether learning is coded in extrinsic or intrinsic reference frames. This dichotomy, however, is challenged by recent findings showing that learning is represented in a mixed reference frame. Overlooked in this framework is how learning appears to consist of multiple processes, such as explicit reaiming and implicit motor adaptation. Therefore, the proposed mixed representation may simply reflect the superposition of explicit and implicit generalization functions, each represented in different reference frames. Here we characterized the individual generalization functions of explicit and implicit learning in relative isolation to determine whether their combination could predict the overall generalization function when both processes are in operation. We modified the form of feedback in a visuomotor rotation task in an attempt to isolate explicit and implicit learning and tested generalization across new limb postures to dissociate the extrinsic/intrinsic representations. We found that the amplitude of explicit generalization was reduced with postural change and was only marginally shifted, resembling an extrinsic representation. In contrast, implicit generalization maintained its amplitude but was significantly shifted, resembling a mixed representation. A linear combination of individual explicit and implicit generalization functions accounted for nearly 85% of the variance associated with the generalization function in a typical visuomotor rotation task, where both processes are in operation. This suggests that each form of learning results from a mixed representation with distinct extrinsic and intrinsic contributions and the combination of these features shapes the generalization pattern observed at novel limb postures. Generalization following learning in visuomotor adaptation tasks can reflect how the brain represents what it learns. In this study, we isolated explicit and implicit forms of learning and showed that they are derived from a mixed reference frame representation with distinct extrinsic and intrinsic contributions. Furthermore, we showed that the overall generalization pattern at novel workspaces is due to the superposition of independent generalization effects developed by explicit and implicit learning processes.