Surprise is a key component of many learning experiences, and yet its precise computational role, and how it changes with age, remain debated. One major challenge is that surprise often occurs jointly with other variables, such as uncertainty and outcome probability. To assess how humans learn from surprising events, and whether aging affects this process, we studied choices while participants learned from bandits with either Gaussian or bi-modal outcome distributions, which decoupled outcome probability, uncertainty, and surprise. A total of 102 participants (51 older, aged 50-73; 51 younger, 19-30 years) chose between three bandits, one of which had a bimodal outcome distribution. Behavioral analyses showed that both age-groups learned the average of the bimodal bandit less well. A trial-by-trial analysis indicated that participants performed choice reversals immediately following large absolute prediction errors, consistent with heightened sensitivity to surprise. This effect was stronger in older adults. Computational models indicated that learning rates in younger as well as older adults were influenced by surprise, rather than uncertainty, but also suggested large interindividual variability in the process underlying learning in our task. Our work bridges between behavioral economics research that has focused on how outcomes with low probability affect choice in older adults, and reinforcement learning work that has investigated age differences in the effects of uncertainty and suggests that older adults overly adapt to surprising events, even when accounting for probability and uncertainty effects.

Humans sometimes have an insight that leads to a sudden and drastic performance improvement on the task they are working on. Sudden strategy adaptations are often linked to insights, considered to be a unique aspect of human cognition tied to complex processes such as creativity or meta-cognitive reasoning. Here, we take a learning perspective and ask whether insight-like behaviour can occur in simple artificial neural networks, even when the models only learn to form input-output associations through gradual gradient descent. We compared learning dynamics in humans and regularised neural networks in a perceptual decision task that included a hidden regularity to solve the task more efficiently. Our results show that only some humans discover this regularity, and that behaviour is marked by a sudden and abrupt strategy switch that reflects an aha-moment. Notably, we find that simple neural networks with a gradual learning rule and a constant learning rate closely mimicked behavioural characteristics of human insight-like switches, exhibiting delay of insight, suddenness and selective occurrence in only some networks. Analyses of network architectures and learning dynamics revealed that insight-like behaviour crucially depended on a regularised gating mechanism and noise added to gradient updates, which allowed the networks to accumulate "silent knowledge" that is initially suppressed by regularised gating. This suggests that insight-like behaviour can arise from gradual learning in simple neural networks, where it reflects the combined influences of noise, gating and regularisation. These results have potential implications for more complex systems, such as the brain, and guide the way for future insight research.

Learning and generalization in spatial domains is often thought to rely on a "cognitive map", representing relationships between spatial locations. Recent research suggests that this same neural machinery is also recruited for reasoning about more abstract, conceptual forms of knowledge. Yet, to what extent do spatial and conceptual reasoning share common computational principles, and what are the implications for behavior? Using a within-subject design we studied how participants used spatial or conceptual distances to generalize and search for correlated rewards in successive multi-armed bandit tasks. Participant behavior indicated sensitivity to both spatial and conceptual distance, and was best captured using a Bayesian model of generalization that formalized distance-dependent generalization and uncertainty-guided exploration as a Gaussian Process regression with a radial basis function kernel. The same Gaussian Process model best captured human search decisions and judgments in both domains, and could simulate realistic learning curves, where we found equivalent levels of generalization in spatial and conceptual tasks. At the same time, we also find characteristic differences between domains. Relative to the spatial domain, participants showed reduced levels of uncertainty-directed exploration and increased levels of random exploration in the conceptual domain. Participants also displayed a one-directional transfer effect, where experience in the spatial task boosted performance in the conceptual task, but not vice versa. While confidence judgments indicated that participants were sensitive to the uncertainty of their knowledge in both tasks, they did not or could not leverage their estimates of uncertainty to guide exploration in the conceptual task. These results support the notion that value-guided learning and generalization recruit cognitive-map dependent computational mechanisms in spatial and conceptual domains. Yet both behavioral and model-based analyses suggest domain specific differences in how these representations map onto actions.

[This corrects the article DOI: 10.1371/journal.pcbi.1008149.].

The activity of neural populations in the brains of humans and animals can exhibit vastly different spatial patterns when faced with different tasks or environmental stimuli. The degrees of similarity between these neural activity patterns in response to different events are used to characterize the representational structure of cognitive states in a neural population. The dominant methods of investigating this similarity structure first estimate neural activity patterns from noisy neural imaging data using linear regression, and then examine the similarity between the estimated patterns. Here, we show that this approach introduces spurious bias structure in the resulting similarity matrix, in particular when applied to fMRI data. This problem is especially severe when the signal-to-noise ratio is low and in cases where experimental conditions cannot be fully randomized in a task. We propose Bayesian Representational Similarity Analysis (BRSA), an alternative method for computing representational similarity, in which we treat the covariance structure of neural activity patterns as a hyper-parameter in a generative model of the neural data. By marginalizing over the unknown activity patterns, we can directly estimate this covariance structure from imaging data. This method offers significant reductions in bias and allows estimation of neural representational similarity with previously unattained levels of precision at low signal-to-noise ratio, without losing the possibility of deriving an interpretable distance measure from the estimated similarity. The method is closely related to Pattern Component Model (PCM), but instead of modeling the estimated neural patterns as in PCM, BRSA models the imaging data directly and is suited for analyzing data in which the order of task conditions is not fully counterbalanced. The probabilistic framework allows for jointly analyzing data from a group of participants. The method can also simultaneously estimate a signal-to-noise ratio map that shows where the learned representational structure is supported more strongly. Both this map and the learned covariance matrix can be used as a structured prior for maximum a posteriori estimation of neural activity patterns, which can be further used for fMRI decoding. Our method therefore paves the way towards a more unified and principled analysis of neural representations underlying fMRI signals. We make our tool freely available in Brain Imaging Analysis Kit (BrainIAK).

In neuroscience, the similarity matrix of neural activity patterns in response to different sensory stimuli or under different cognitive states reflects the structure of neural representational space. Existing methods derive point estimations of neural activity patterns from noisy neural imaging data, and the similarity is calculated from these point estimations. We show that this approach translates structured noise from estimated patterns into spurious bias structure in the resulting similarity matrix, which is especially severe when signal-to-noise ratio is low and experimental conditions cannot be fully randomized in a cognitive task. We propose an alternative Bayesian framework for computing representational similarity in which we treat the covariance structure of neural activity patterns as a hyper-parameter in a generative model of the neural data, and directly estimate this covariance structure from imaging data while marginalizing over the unknown activity patterns. Converting the estimated covariance structure into a correlation matrix offers a much less biased estimate of neural representational similarity. Our method can also simultaneously estimate a signal-to-noise map that informs where the learned representational structure is supported more strongly, and the learned covariance matrix can be used as a structured prior to constrain Bayesian estimation of neural activity patterns. Our code is freely available in Brain Imaging Analysis Kit (Brainiak) (https://github.com/IntelPNI/brainiak), a python toolkit for brain imaging analysis.