The relation between the hippocampal spatial code and internal
representations of goals has been a topic of discussions since the
discovery of place cells. Today most neuroscientists agree that the firing
of hippocampal place cells in an alert animal encodes for the generalized
context in which the subject is situated rather than the spatial location
only. This view is consistent with the so-called remapping phenomena: the
hippocampus may exhibit very different spatial codes for the same
environment in behavioral episodes that differ by their paradigms, and
even in different phases of the same paradigm performed repeatedly. In
particular, the generalized context encoded by place cells could include
the prospect of reward.
The evidence presented by Hok et al. supports this position. When the
firing patterns are decomposed into different phases of the paradigm (Fig.
1D), two distinct spatial maps become apparent: the foraging map (Fig. 1D,
right panel) and the goal-zone map (Fig. 1D, small circle in the middle
panel), rather than one persistent map with excess firing at the goal
location (Fig. 1D, left panel). In fact, the foraging map shows no surplus
activity at the goal-zone. We surmise that the goal-zone map only exists
for a short temporal window as a global hippocampal state, when the rat
expects a reward in the goal-zone, but not during the larger time interval
labeled by the authors as "navigation". Given this assumption, this map
would be limited in space simply because it is limited in time. The goal-
zone map is only defined and makes sense for the small circle, exactly as
the foraging map is defined and makes sense for the large circle. The fact
that most cells show up on the goal-zone map follows from the general
observation that most place cells active on one map typically fire on
another map in the same environment.
This parsimonious explanation invites the hypothesis that the
excessive goal-zone firing could play a functional role in learning how to
navigate to the goal-zone. In the model of hippocampal navigation we
recently proposed (Samsonovich & Ascoli, 2005), any CA1 cell with a
place field in a familiar environment is potentially available as a goal
cell. The model predicts that place cells that fire on the way to the
(actual or potential) goal get reactivated when the goal location is
reached, enabling reinforcement learning. Available data indicate that
this necessary reactivation occurs during sharp waves. The finding of Hok
et al. suggests that reactivation could happen during theta, possibly
under circumstances specific to this paradigm. Interestingly, the authors
noticed switching from theta-I to theta-II in the goal-zone, which in
agreement with the observed synchrony of firing might be associated with
sharp waves.
As a key component of the spatial memory system, the hippocampus is
frequently implicated in spatial navigation, and yet direct evidence of
this function is missing. What mechanism (if any) connects the hippocampal
place code with the ability to navigate towards a goal? In our model, the
same neural mechanism, based on reinforcement learning, linked spatial and
mnemonic functions by implementing episodic memory retrieval as contextual
pathfinding (Samsonovich & Ascoli 2005). The results of Hok et al.
could thus extend to human episodic memory, implying that most hippocampal
pointers, rather than only one or few, should activate when subjects
actively engage in context retrieval.
Samsonovich AV, Ascoli GA ( 2005) A simple neural network model of
the hippocampus suggesting its pathfinding role in episodic memory
retrieval. Learning & Memory 12(2):193–208.