ABSTRACT:The oscillatory interference model Hippocampus 17:801-812) explains the generation of spatially stable, regular firing patterns by medial entorhinal cortical (mEC) grid cells in terms of the interference between velocity-controlled oscillators (VCOs) with different preferred directions. This model predicts specific relationships between the intrinsic firing frequency and spatial scale of grid cell firing, the EEG theta frequency, and running speed (Burgess, 2008). Here, we use spectral analyses of EEG and of spike autocorrelograms to estimate the intrinsic firing frequency of grid cells, and the concurrent theta frequency, in mEC Layer II in freely moving rats. The intrinsic firing frequency of grid cells increased with running speed and decreased with grid scale, according to the quantitative prediction of the model. Similarly, theta frequency increased with running speed, which was also predicted by the model. An alternative Moiré interference model (Blair et al., 2007) predicts a direction-dependent variation in intrinsic firing frequency, which was not found. Our results suggest that interference between VCOs generates the spatial firing patterns of entorhinal grid cells according to the oscillatory interference model. They also provide specific constraints on this model of grid cell firing and have more general implications for viewing neuronal processing in terms of interfering oscillatory processes. V
The hippocampal formation (HF) plays a key role in novelty detection, but the mechanisms remain unknown. Novelty detection aids the encoding of new information into memory-a process thought to depend on the HF and to be modulated by the theta rhythm of EEG. We examined EEG recorded in the HF of rats foraging for food within a novel environment, as it became familiar over the next five days, and in two more novel environments unexpectedly experienced in trials interspersed with familiar trials over three further days. We found that environmental novelty produces a sharp reduction in the theta frequency of foraging rats, that this reduction is greater for an unexpected environment than for a completely novel one, and that it slowly disappears with increasing familiarity. These results do not reflect changes in running speed and suggest that the septo-hippocampal system signals unexpected environmental change via a reduction in theta frequency. In addition, they provide evidence in support of a cholinergically mediated mechanism for novelty detection, have important implications for our understanding of oscillatory coding within memory and for the interpretation of event-related potentials, and provide indirect support for the oscillatory interference model of grid cell firing in medial entorhinal cortex.
Hippocampal processing is strongly implicated in both spatial cognition and anxiety and is temporally organized by the theta rhythm. However, there has been little attempt to understand how each type of processing relates to the other in behaving animals, despite their common substrate. In freely moving rats, there is a broadly linear relationship between hippocampal theta frequency and running speed over the normal range of speeds used during foraging. A recent model predicts that spatial-translation-related and arousal/anxietyrelated mechanisms of hippocampal theta generation underlie dissociable aspects of the theta frequency-running speed relationship (the slope and intercept, respectively). Here we provide the first confirmatory evidence: environmental novelty decreases slope, whereas anxiolytic drugs reduce intercept. Variation in slope predicted changes in spatial representation by CA1 place cells and noveltyresponsive behavior. Variation in intercept predicted anxiety-like behavior. Our findings isolate and doubly dissociate two components of theta generation that operate in parallel in behaving animals and link them to anxiolytic drug action, novelty, and the metric for self-motion.
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