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.
Pigeons are classic model animals to study perceptual category learning. To achieve a deeper understanding of the cognitive mechanisms of categorization, a careful consideration of the employed stimulus material and a thorough analysis of the choice behavior is mandatory. In the present study, we combined the use of “virtual phylogenesis”, an evolutionary algorithm to generate artificial yet naturalistic stimuli termed digital embryos and a machine learning approach on the pigeons’ pecking responses to gain insight into the underlying categorization strategies of the animals. In a forced-choice procedure, pigeons learned to categorize these stimuli and transferred their knowledge successfully to novel exemplars. We used peck tracking to identify where on the stimulus the animals pecked and further investigated whether this behavior was indicative of the pigeon’s choice. Going beyond the classical analysis of the binary choice, we were able to predict the presented stimulus class based on pecking location using a k-nearest neighbor classifier, indicating that pecks are related to features of interest. By analyzing error trials with this approach, we further identified potential strategies of the pigeons to discriminate between stimulus classes. These strategies remained stable during category transfer, but differed between individuals indicating that categorization learning is not limited to a single learning strategy.
Pigeons are classic model animals to study perceptual category learning. A theoretical understanding of the cognitive mechanisms of categorization requires a careful consideration of the employed stimulus material. Optimally, stimuli should not consist of real-world objects that might be associated with prior experience. The number of exemplars should be theoretically infinite and easy to produce. In addition, the experimenter should have the freedom to produce 2D- and 3D-versions of the stimuli and, finally, the stimulus set should provide the opportunity to identify the diagnostic elements that the animals use. To this end, we used the approach of "virtual phylogenesis" of "digital embryos" to produce two stimulus sets of objects that meet these criteria. In our experiment pigeons learned to categorize these stimuli in a forced-choice procedure. In addition, we used peck tracking to identify where on the stimulus the animals pecked to signal their choice. Pigeons learned the task and transferred successfully to novel exemplars. Using a k-nearest neighbor classifier, we were able to predict the presented stimulus class based on pecking location indicating that pecks are related to features of interest. We further identified potential strategies of the pigeons through this approach, namely that they were either learning one or two categories to discriminate between stimulus classes. These strategies remained stable during category transfer, but differed between individuals indicating that categorization learning is not limited to a single learning strategy.
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