Thus far, we have focused on behavioral, lesion, and fMRI studies, which argue against allocentric navigational strategies depending on a single brain region (Figure 3A) and as decomposable into contributions from individual brain regions (Figure 3B). One might argue, as others have (O’Keefe and Nadel, 1978; Redish, 1999), however, that place cells, present in the rodent, monkey, and human hippocampus (O’Keefe and Dostrovsky, 1971; Ekstrom et al., 2003; Hori et al., 2003), are the neural instantiation of an allocentric representation, or cognitive map. While place cells do have many features similar to what one might expect in a neural systems that code spatial environments in a map-like fashion, there are other important features of place cells that are decidedly not map-like. Place cells in the rodent and human hippocampus remap based on egocentric direction (Markus et al., 1995; Miller et al., 2013), are sensitive to goal and other temporal variables (Gothard et al., 2001; Hollup et al., 2001; Ekstrom et al., 2003; Bahar et al., 2011), and remap with subtle changes to the spatial geometry of the environment (Leutgeb et al., 2005; Wills et al., 2005). Indeed, recent theoretical models of the cognitive map now suggest that time and geometry less variant spatial coding mechanisms possibly resides outside of the hippocampus (Buzsaki, 2006; Buzsaki and Moser, 2013). Grid cells, neurons in enthorhinal cortex that fire in a regularly spaced fashion as the rat explores a spatial environment (Fyhn et al., 2004; Jacobs et al., 2013), may be a better candidate for the neural basis of allocentric representation (Buzsaki and Moser, 2013). Yet lesions of entorhinal cortex, at least in rodents, do not abolish place cell firing in the CA3 subfield of the hippocampus (Lu et al., 2013) and impair, but do not abolish, the place code in CA1 (Brun et al., 2008). While many details of entorhinal–hippocampal neural interactions remain to be established, grid cells do not contribute in a clear or modular fashion to place coding in the hippocampus, at least based on what the above-mentioned studies have determined so far in the rat. Furthermore, in addition to grid cells, entorhinal cortex cells also respond to egocentric direction (Sargolini et al., 2006), suggesting this area may not be specialized for allocentric computations alone. In addition, consistent with what we have argued here, it is clear that other areas, like prefrontal and retrosplenial cortex, also contribute critically, via oscillatory synchrony, to spatial coding in the hippocampus (Benchenane et al., 2010; Battaglia et al., 2011; Fujisawa and Buzsaki, 2011). Thus, although many aspects of the hippocampal neural code would appear sufficient to support an allocentric representation, the neural code itself is not map-like and depends, at least in part, on coordinated input and activity from other brain structures.