Although hippocampal grid cells are thought to be crucial for spatial navigation, their computational purpose remains disputed. Recently, they were proposed to represent spatial transitions and convey this knowledge downstream to place cells. However, a single scale of transitions is insufficient to plan long goal-directed sequences in behaviorally acceptable time. Here, a scale-space data structure is suggested to optimally accelerate retrievals from transition systems, called transition scale-space (TSS). Remaining exclusively on an algorithmic level, the scale increment is proved to be ideally 2 for biologically plausible receptive fields. It is then argued that temporal buffering is necessary to learn the scale-space online. Next, two modes for retrieval of sequences from the TSS are presented: top down and bottom up. The two modes are evaluated in symbolic simulations (i.e., without biologically plausible spiking neurons). Additionally, a TSS is used for short-cut discovery in a simulated Morris water maze. Finally, the results are discussed in depth with respect to biological plausibility, and several testable predictions are derived. Moreover, relations to other grid cell models, multiresolution path planning, and scale-space theory are highlighted. Summarized, reward-free transition encoding is shown here, in a theoretical model, to be compatible with the observed discretization along the dorso-ventral axis of the medial entorhinal cortex. Because the theoretical model generalizes beyond navigation, the TSS is suggested to be a general-purpose cortical data structure for fast retrieval of sequences and relational knowledge. Source code for all simulations presented in this paper can be found at https://github.com/rochus/transitionscalespace .

Grid cells of the rodent entorhinal cortex are essential for spatial navigation. Although their function is commonly believed to be either path integration or localization, the origin or purpose of their hexagonal firing fields remains disputed. Here they are proposed to arise as an optimal encoding of transitions in sequences. First, storage requirements for transitions in general episodic sequences are examined using propositional logic and graph theory. Subsequently, transitions in complete metric spaces are considered under the assumption of an ideal sampling of an input space. It is shown that memory capacity of neurons that have to encode multiple feasible spatial transitions is maximized by a hexagonal pattern. Grid cells are proposed to encode spatial transitions in spatiotemporal sequences, with the entorhinal-hippocampal loop forming a multitransition system.