“… θ : Grid cells have a discrete organization into grid modules in MEC, where up to 10 grid modules, each with a single orientation, θ , are estimated to exist in the rat MEC (Stensola et al, ). Border cells (Barry et al, ; Krupic, Bauza, Burton, & O'Keefe, ; Lever, Burton, Jeewajee, O'Keefe, & Burgess, ; Solstad, Boccara, Kropff, Moser, & Moser, ) in MEC are active when the rat is at the boundary of an enclosure. Along with evidence that grid cells can show different orientations in a single environment if there are distorted boundaries (Krupic, Bauza, Burton, Barry, & O'Keefe, ), this suggests boundary cells may orient grid cells.…”