What about timing? As the rat passes through a place field, the associated place cell fires earlier and earlier with respect to the cycle of the background theta wave. As the rat crosses from one place field into another, the very early firing of the first place cell occurs close in time with the late firing of the next place cell. Their near-coincident firings cause the synapse between them to strengthen, and this coupling of the place cells ingrains the rat’s trajectory into the brain (strengthening of synapses occurs only when two neurons fire within tens of milliseconds of each other.)
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