Multilevel structure in behaviour and in the brain: a model of Fuster's hierarchy

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Abstract
A basic question, intimately tied to the problem of action selection, is that of how actions are assembled into organized sequences. Theories of routine sequential behaviour have long acknowledged that it must rely not only on environmental cues but also on some internal representation of temporal or task context. It is assumed, in most theories, that such internal representations must be organized into a strict hierarchy, mirroring the hierarchical structure of naturalistic sequential behaviour. This article reviews an alternative computational account, which asserts that the representations underlying naturalistic sequential behaviour need not, and arguably cannot, assume a strictly hierarchical form. One apparent liability of this theory is that it seems to contradict neuroscientific evidence indicating that different levels of sequential structure in behaviour are represented at different levels in a hierarchy of cortical areas. New simulations, reported here, show not only that the original computational account can be reconciled with this alignment between behavioural and neural organization, but also that it gives rise to a novel explanation for how this alignment might develop through learning.