Integration of the sensory inputs to place cells: what, where, why, and how?

The hippocampal place cells are a highly multimodal class of neurons, receiving information from many different sensory sources to correctly localize their firing to restricted regions of an environment. Evidence suggests that the sensory information is processed upstream of the hippocampus, to extract both angular and linear metric information, and also contextual information. These various kinds of information need to be integrated for coherent firing fields to be generated, and the present article reviews recent evidence concerning how this occurs. It is concluded that there is a functional dissociation of the cortical inputs, with one class of incoming information comprising purely metric information concerning distance and orientation, probably routed via the grid cells and head direction cells. The other class of information is much more heterogeneous and serves, at least in part, to contextualize the spatial inputs so as to provide a unique representation of the place the animal is in. Evidence from remapping studies suggests that the metric and contextual inputs interact upstream of the place cells, perhaps in entorhinal cortex. A full understanding of the generation of the hippocampal place representation will require elucidation of the representational functions of the afferent cortical areas.