The neuroscience of perceptual categorization in pigeons: A mechanistic hypothesis.

We are surrounded by an endless variation of objects. The ability to categorize these objects represents a core cognitive competence of humans and possibly all vertebrates. Research on category learning in nonhuman animals started with the seminal studies of Richard Herrnstein on the category "human" in pigeons. Since then, we have learned that pigeons are able to categorize a large number of stimulus sets, ranging from Cubist paintings to English orthography. Strangely, this prolific field has largely neglected to also study the avian neurobiology of categorization. Here, we present a hypothesis that combines experimental results and theories from categorization research in pigeons with neurobiological insights on visual processing and dopamine-mediated learning in primates. We conclude that in both fields, similar conclusions on the mechanisms of perceptual categorization have been drawn, despite very little cross-reference or communication between these two areas to date. We hypothesize that perceptual categorization is a two-component process in which stimulus features are first rapidly extracted in a feed-forward process, thereby enabling a fast subdivision along multiple category borders. In primates this seems to happen in the inferotemporal cortex, while pigeons may primarily use a cluster of associative visual forebrain areas. The second process rests on dopaminergic error-prediction learning that enables prefrontal areas to connect top down the relevant visual category dimension to the appropriate action dimension.