Understanding how grid cells perform path integration calculations remains a fundamental problem. In this paper, we conduct theoretical analysis of a general representation model of path integration by grid cells, where the 2D self-position is encoded as a higher dimensional vector, and the 2D self-motion is represented by a general transformation of the vector. We identify two conditions on the transformation. One is a group representation condition that is necessary for path integration. The other is an isotropic scaling condition that ensures locally conformal embedding, so that the error in the vector representation translates proportionally to the error in the 2D self-position. Then we investigate the simplest transformation, i.e., the linear transformation, uncover its explicit algebraic and geometric structure as matrix Lie group of rotation, and establish the connection between the isotropic scaling condition and hexagon grid patterns of grid cells under the linear transformation. Finally, with our optimization-based approach, we manage to learn hexagon grid patterns that share similar properties of the grid cells in the rodent brain. The learned model is capable of accurate long distance path integration.
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