Neural circuits of hierarchical visuo-spatial sequence processing.

Sequence processing has been investigated in a number of studies using serial reaction time tasks or simple artificial grammar tasks. Little, however, is known about higher-order sequence processing entailing the hierarchical organization of events. Here, we manipulated the regularities within sequentially occurring, non-linguistic visual symbols by applying two types of prediction rules. In one rule (the adjacent dependency rule), the sequences consisted of alternating items from two different categories. In the second rule (the hierarchical dependency rule), a hierarchical structure was generated using the same set of item types. Thus, predictions about non-adjacent elements were required for the latter rule. Functional Magnetic Resonance Imaging (fMRI) was used to investigate the neural correlates of the application of the two prediction rules. We found that the hierarchical dependency rule correlated with activity in the pre-supplementary motor area, and the head of the caudate nucleus. In addition, in a hypothesis-driven ROI analysis in Broca's area (BA 44), we found a significantly higher hemodynamic response to the hierarchical dependency rule than to the adjacent dependency rule. These results suggest that this neural network supports hierarchical sequencing, possibly contributing to the integration of sequential elements into higher-order structural events. Importantly, the findings suggest that Broca's area is also engaged in hierarchical sequencing in domains other than language.