in the environment, the place field. They are described as having a low resting firing rate (<1 Hz) when a rat is not in its place field, but a particularly high firing rate, which can be over 100 Hz in some cases, within the place field. Additionally, O'Keefe described 6 special cells, which he called misplace units, which also fire only in a particular place, but only when the rat performed an additional behaviour, such as sniffing, which was often correlated with the presence of a novel stimulus, or the absence of an expected stimulus. The findings ultimately supported the cognitive map theory, the idea that the hippocampus hold a spatial representation, a cognitive map of the environment. There has been much debate as to whether hippocampal place cells function depends on landmarks in the environment, on environmental boundaries, or on an interaction between the two. Additionally, not all place cells rely on the same external cues. One important distinction in cues is local and distal, where local cues appear in the immediate vicinity of a subject, whereas distal cues are far away, and act more like landmarks. Individual place cells have been shown to follow either or rely on both. Additionally, the cues on which the place cells rely may depend on previous experience of the subject and the saliency of the cue. There has also been much debate as to whether hippocampal pyramidal cells truly encode non-spatial information as well as spatial information. According to the cognitive map theory, the hippocampus's primary role is to store spatial information through place cells and the hippocampus was biologically designed to provide a subject with spatial information. Recent findings, such as a study showing that place cells respond to non-spatial dimensions, such as sound frequency, disagree with the cognitive map theory. Instead, they support a new theory saying that the hippocampus has a more general function encoding continuous variables, and location just happens to be one of those variables. This fits in with the idea that the hippocampus has a predictive function. Relationship to grid cells It has been proposed that place cells are derivatives of grid cells, pyramidal cells in the entorhinal cortex. This theory suggests that the place fields of the place cells are a combination of several grid cells, which have hexagonal grid-like patterns of activity. The theory has been supported by computational models. The relation may arise