Desiderata of evidence for representation in neuroscience

This paper develops a systematic framework for the evidence neuroscientists use to establish whether a neural response represents a feature. Researchers try to establish that the neural response is (1) sensitive and (2) specific to the feature, (3) invariant to other features, and (4) functional, which means that it is used downstream in the brain. We formalize these desiderata in information-theoretic terms. This formalism allows us to precisely state the desiderata while unifying the different analysis methods used in neuroscience under one framework. We discuss how common methods such as correlational analyses, decoding and encoding models, representational similarity analysis, and tests of statistical dependence are used to evaluate the desiderata. In doing so, we provide a common terminology to researchers that helps to clarify disagreements, to compare and integrate results across studies and research groups, and to identify when evidence might be missing and when evidence for some representational conclusion is strong. We illustrate the framework with several canonical examples, including the representation of orientation, numerosity, faces, and spatial location. We end by discussing how the framework can be extended to cover models of the neural code, multi-stage models, and other domains.