Category knowledge can be explicit, yet not conform to a perfect rule. For example, a child may acquire the rule "If it has wings, then it is a bird," but then must account for exceptions to this rule, such as bats. The current study explored the neurobiological basis of rule-plus-exception learning by using quantitative predictions from a category learning model, SUSTAIN, to analyze behavioral and functional magnetic resonance imaging (fMRI) data. SUSTAIN predicts that exceptions require formation of specialized representations to distinguish exceptions from rule-following items in memory. By incorporating quantitative trial-by-trial predictions from SUSTAIN directly into fMRI analyses, we observed medial temporal lobe (MTL) activation consistent with 2 predicted psychological processes that enable exception learning: item recognition and error correction. SUSTAIN explains how these processes vary in the MTL across learning trials as category knowledge is acquired. Importantly, MTL engagement during exception learning was not captured by an alternate exemplar-based model of category learning or by standard contrasts comparing exception and rule-following items. The current findings thus provide a well-specified theory for the role of the MTL in category learning, where the MTL plays an important role in forming specialized category representations appropriate for the learning context.
Multi-voxel pattern analysis (MVPA) has led to major changes in how fMRI data are analyzed and interpreted. Many studies now report both MVPA results and results from standard univariate voxel-wise analysis, often with the goal of drawing different conclusions from each. Because MVPA results can be sensitive to latent multidimensional representations and processes whereas univariate voxel-wise analysis cannot, one conclusion that is often drawn when MVPA and univariate results differ is that the activation patterns underlying MVPA results contain a multidimensional code. In the current study, we conducted simulations to formally test this assumption. Our findings reveal that MVPA tests are sensitive to the magnitude of voxel-level variability in the effect of a condition within subjects, even when the same linear relationship is coded in all voxels. We also find that MVPA is insensitive to subject-level variability in mean activation across an ROI, which is the primary variance component of interest in many standard univariate tests. Together, these results illustrate that differences between MVPA and univariate tests do not afford conclusions about the nature or dimensionality of the neural code. Instead, targeted tests of the informational content and/or dimensionality of activation patterns are critical for drawing strong conclusions about the representational codes that are indicated by significant MVPA results.
Recently, there has been a dramatic increase in the number of functional magnetic resonance imaging studies seeking to answer questions about how the brain represents information. Representational questions are of particular importance in connecting neuroscientific and cognitive levels of analysis because it is at the representational level that many formal models of cognition make distinct predictions. This review discusses techniques for univariate, adaptation, and multivoxel analysis, and how they have been used to answer questions about content specificity in different regions of the brain, how this content is organized, and how representations are shaped by and contribute to cognitive processes. Each of the analysis techniques makes different assumptions about the underlying neural code and thus differ in how they can be applied to specific questions. We also discuss the many pitfalls of representational analysis, from the flexibility in data analysis pipelines to emergent nonrepresentational relationships that can arise between stimuli in a task.
Familiarity, or memory strength, is a central construct in models of cognition. In previous categorization and long-term memory research, correlations have been found between psychological measures of memory strength and activation in the medial temporal lobes (MTLs), which suggests a common neural locus for memory strength. However, activation alone is insufficient for determining whether the same mechanisms underlie neural function across domains. Guided by mathematical models of categorization and long-term memory, we develop a theory and a method to test whether memory strength arises from the global similarity among neural representations. In human subjects, we find significant correlations between global similarity among activation patterns in the MTLs and both subsequent memory confidence in a recognition memory task and model-based measures of memory strength in a category learning task. Our work bridges formal cognitive theories and neuroscientific models by illustrating that the same global similarity computations underlie processing in multiple cognitive domains. Moreover, by establishing a link between neural similarity and psychological memory strength, our findings suggest that there may be an isomorphism between psychological and neural representational spaces that can be exploited to test cognitive theories at both the neural and behavioral levels.
Category learning is a complex phenomenon that engages multiple cognitive processes, many of which occur simultaneously and unfold dynamically over time. For example, as people encounter objects in the world, they simultaneously engage processes to determine their fit with current knowledge structures, gather new information about the objects, and adjust their representations to support behavior in future encounters. Many techniques that are available to understand the neural basis of category learning assume that the multiple processes that subserve it can be neatly separated between different trials of an experiment. Model-based functional magnetic resonance imaging offers a promising tool to separate multiple, simultaneously occurring processes and bring the analysis of neuroimaging data more in line with category learning’s dynamic and multifaceted nature. We use model-based imaging to explore the neural basis of recognition and entropy signals in the medial temporal lobe and striatum that are engaged while participants learn to categorize novel stimuli. Consistent with theories suggesting a role for the anterior hippocampus and ventral striatum in motivated learning in response to uncertainty, we find that activation in both regions correlates with a model-based measure of entropy. Simultaneously, separate subregions of the hippocampus and striatum exhibit activation correlated with a model-based recognition strength measure. Our results suggest that model-based analyses are exceptionally useful for extracting information about cognitive processes from neuroimaging data. Models provide a basis for identifying the multiple neural processes that contribute to behavior, and neuroimaging data can provide a powerful test bed for constraining and testing model predictions.
We suggest that human category formation relies on contrastive learning mechanisms that seek to reduce prediction error. In keeping with this view, manipulating category contrast leads to systematic distortions in people's memory for category information. Simply by changing the basis of comparison (i.e., the available response options), we can systematically distort people's perceptions of novel, energy-source, and political categories. Our proposal explains perceived variations in category members' typicality, including cases in which average items are judged as highly typical and cases in which extreme or ideal members are judged as highly typical of the category. Although straightforward, our account spans findings from studies in goal-derived, cross-cultural, and object-based categorization and suggests ways in which society's perception of key issues is distorted by political discourse.
How categories are represented continues to be hotly debated across neuroscience and psychology. One topic that is central to cognitive research on category representation but underexplored in neurobiological research concerns the internal structure of categories. Internal structure refers to how the natural variability between-category members is coded so that we are able to determine which members are more typical or better examples of their category. Psychological categorization models offer tools for predicting internal structure and suggest that perceptions of typicality arise from similarities between the representations of category members in a psychological space. Inspired by these models, we develop a neural typicality measure that allows us to measure which category members elicit patterns of activation that are similar to other members of their category and are thus more central in a neural space. Using an artificial categorization task, we test how psychological and physical typicality contribute to neural typicality, and find that neural typicality in occipital and temporal regions is significantly correlated with subjects' perceptions of typicality. The results reveal a convergence between psychological and neural category representations and suggest that our neural typicality measure is a useful tool for connecting psychological and neural measures of internal category structure.
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