Despite the centrality of the notion of representation to its explanations, neuroscience lacks a unified framework for the concepts used to characterize representation, leading to disparate use of terminology and measures associated with representation. To offer clarification, we propose a core set of conceptual dimensions that characterize representations in neuroscience. These dimensions describe relations between a neural response, features that may be represented, and downstream effects of the neural response. A neural response may be shown to be sensitive and specific to a feature, invariant to other features, and functional, which means that it is used downstream in the brain. We use information-theoretic measures to introduce these conceptual dimensions unambiguously and explain how data analysis methods such as correlational analyses, decoding and encoding models, representational similarity analysis, and tests of statistical dependence or adaptation relate to our framework. We consider several canonical examples, including the representation of orientation, numerosity, and spatial location, which illustrate how the evidence put forth in support or criticism of representational conclusions is systematized by our framework. By offering a unified conceptual framework we hope to aid the comparison and integration of results across studies and research groups and to help determine when evidence for a representational conclusion is strong.

Perceptual decision-making is often conceptualized as the process of comparing an internal decision variable to a categorical boundary or criterion. How the mind sets such a criterion has been studied from at least two perspectives. One idea is that the criterion is a fixed quantity. In work on subjective phenomenology, the notion of a fixed criterion has been proposed to explain a phenomenon called "subjective inflation"-a form of metacognitive mismatch in which observers overestimate the quality of their sensory representation in the periphery or at unattended locations. A contrasting view emerging from studies of perceptual decision-making is that the criterion adjusts to the level sensory uncertainty and is thus sensitive to variations in attention. Here, we mathematically demonstrate that previous empirical findings supporting subjective inflation are consistent with either a fixed or a flexible decision criterion. We further lay out specific task properties that are necessary to make inferences about the flexibility of the criterion: (i) a clear mapping from decision variable space to stimulus feature space and (ii) an incentive for observers to adjust their decision criterion as uncertainty changes. Recent work satisfying these requirements has demonstrated that decision criteria flexibly adjust according to uncertainty. We conclude that the fixed-criterion model of subjective inflation is poorly tenable.

The study of the brain's representations of uncertainty is a central topic in neuroscience. Unlike most quantities of which the neural representation is studied, uncertainty is a property of an observer's beliefs about the world, which poses specific methodological challenges. We analyze how the literature on the neural representations of uncertainty addresses those challenges and distinguish between 'code-driven' and 'correlational' approaches. Code-driven approaches make assumptions about the neural code for representing world states and the associated uncertainty. By contrast, correlational approaches search for relationships between uncertainty and neural activity without constraints on the neural representation of the world state that this uncertainty accompanies. To compare these two approaches, we apply several criteria for neural representations: sensitivity, specificity, invariance and functionality. Our analysis reveals that the two approaches lead to different but complementary findings, shaping new research questions and guiding future experiments.

Prediction of future sensory input based on past sensory information is essential for organisms to effectively adapt their behavior in dynamic environments. Humans successfully predict future stimuli in various natural settings. Yet, it remains elusive how the brain achieves effective prediction despite enormous variations in sensory input rate, which directly affect how fast sensory information can accumulate. We presented participants with acoustic sequences capturing temporal statistical regularities prevalent in nature and investigated neural mechanisms underlying predictive computation using MEG. By parametrically manipulating sequence presentation speed, we tested two hypotheses: neural prediction relies on integrating past sensory information over fixed time periods or fixed amounts of information. We demonstrate that across halved and doubled presentation speeds, predictive information in neural activity stems from integration over fixed amounts of information. Our findings reveal the neural mechanisms enabling humans to robustly predict dynamic stimuli in natural environments despite large sensory input rate variations.

The spatial distribution of visual items allows us to infer the presence of latent causes in the world. For instance, a spatial cluster of ants allows us to infer the presence of a common food source. However, optimal inference requires the integration of a computationally intractable number of world states in real world situations. For example, optimal inference about whether a common cause exists based on N spatially distributed visual items requires marginalizing over both the location of the latent cause and 2N possible affiliation patterns (where each item may be affiliated or non-affiliated with the latent cause). How might the brain approximate this inference? We show that subject behaviour deviates qualitatively from Bayes-optimal, in particular showing an unexpected positive effect of N (the number of visual items) on the false-alarm rate. We propose several "point-estimating" observer models that fit subject behaviour better than the Bayesian model. They each avoid a costly computational marginalization over at least one of the variables of the generative model by "committing" to a point estimate of at least one of the two generative model variables. These findings suggest that the brain may implement partially committal variants of Bayesian models when detecting latent causes based on complex real world data.

Forming valid predictions about the environment is crucial to survival. However, whether humans are able to form valid predictions about natural stimuli based on their temporal statistical regularities remains unknown. Here, we presented subjects with tone sequences with pitch fluctuations that, over time, capture long-range temporal dependence structures prevalent in natural stimuli. We found that subjects were able to exploit such naturalistic statistical regularities to make valid predictions about upcoming items in a sequence. Magnetoencephalography (MEG) recordings revealed that slow, arrhythmic cortical dynamics tracked the evolving pitch sequence over time such that neural activity at a given moment was influenced by the pitch of up to seven previous tones. Importantly, such history integration contained in neural activity predicted the expected pitch of the upcoming tone, providing a concrete computational mechanism for prediction. These results establish humans' ability to make valid predictions based on temporal regularities inherent in naturalistic stimuli and further reveal the neural mechanisms underlying such predictive computation.SIGNIFICANCE STATEMENT A fundamental question in neuroscience is how the brain predicts upcoming events in the environment. To date, this question has primarily been addressed in experiments using relatively simple stimulus sequences. Here, we studied predictive processing in the human brain using auditory tone sequences that exhibit temporal statistical regularities similar to those found in natural stimuli. We observed that humans are able to form valid predictions based on such complex temporal statistical regularities. We further show that neural response to a given tone in the sequence reflects integration over the preceding tone sequence and that this history dependence forms the foundation for prediction. These findings deepen our understanding of how humans form predictions in an ecologically valid environment.