example, faces versus natural scenes (Kreiman, Koch, Fried, 2000) and

when the participant viewed or recalled specific video clips (Gelbard-Sagiv,

Mukamel, Harel, Malach, & Fried, 2008). Hippocampal neurons in
monkeys have been shown to fire preferentially when specific objects
(Naya & Suzuki, 2011) or visual scenes were presented to the monkey, and
the firing rates of these neurons showed modulations in activity that
correlated with the monkeys’ ability to learn to move their eyes to a
particular place in the scene to receive a reward (Wirth et al., 2003). Studies
in rats have also shown that hippocampal neurons sometimes fire
preferentially for specific olfactory stimuli as well as combinations of these
stimuli and the places where they occur, suggesting that hippocampal
neurons represent important stimuli in the context in which they are
remembered (Wood, Dudchenko, & Eichenbaum, 1999).
In addition to this activity related to memory for items, neurons with
spatial correlates have been identified in the hippocampal memory system
of several species of mammals, including humans (O’Keefe & Dostrovsky,
1971; Derdikman & Moser, 2010; also see Chapter 45). In particular, many
neurons in the rat hippocampus, referred to as “place cells,” have clearly
defined spatial receptive fields, or “place fields” (Fig. 48.8A). Activity of
these cells reflects the constellation of auditory, olfactory, tactile, and visual
cues that define the external world as well as the self-motion cues that track
the rat’s path through that space. Yet in addition to these clear spatial
correlates, activity of these cells also reflects factors such as goals and
expectations, and a common view is that the type of spatial memory
reflected by place cells in rats shares many commonalities with examples of
memory in humans such as recall. Further, cells with location or direction
correlates have been found in several regions interconnected with the
hippocampus, leading to an understanding of how mental maps might be
constructed in the brain. An important advancement in that goal was the
identification of “grid cells” in a region of the medial entorhinal cortex, an
area reciprocally connected with the hippocampus (Hafting, Fyhn, Molden,
Moser, & Moser, 2005). The activity of one of these cells tiles a recording
chamber in a geometrically regular grid pattern, and the overlapping
receptive fields of the population of grid cells in a rat’s brain are thought to
be an essential component of the rat’s mental map (Fig. 48.8B). An
important question is how the circuitry of the hippocampus and entorhinal
cortex (and other connected areas) enables the transformation of spatial