Navigation requires many different processes, such as generation of body
movements, recognition of landmarks, sense of direction, memory for
routes and memory for survey-type knowledge. Thus disentangling the
neural correlates of navigation will not be easy. Before considering the
uses of VR in this pursuit, we briefly summarize the background to this
area.
Electrophysiological recording in awake behaving animals has revealed
several types of representation of spatial location in the brain. Neurons in
posterior parietal areas have been found to represent the location of
stimuli or actions in reference frames related to parts of the body
(‘egocentric’ reference frames). For example, the firing rates of neurons in
the posterior parietal cortex of monkeys may reflect the location of a
stimulus relative to its eye or head or body. They also appear to encode
the information necessary to translate between reference frames (e.g.,
given a location relative to the eye, computing that location relative to the
head. See Andersen et al. 1985). Of particular interest with respect to
navigation, ‘place cells’ in the rat hippocampus represent the rat’s current
location with respect to its environment (an ‘allocentric’ reference frame
(e.g., O’Keefe and Nadel 1978). These cells become active whenever the
rat is in a particular portion of an open environment, independent of the
rat’s orientation. The complementary representation, of heading direction
independent of location (‘head-direction cells’), was found in an area close
to the hippocampus, the presubiculum.
Consistent with the finding of place cells, and the idea that the
hippocampus maintains a survey-type representation or ‘cognitive map’ of
the environment (O’Keefe and Nadel 1978), lesions to the hippocampus
impair a rat’s ability to navigate to the hidden platform in the Morris water
maze (Morris et al. 1982).
The egocentric parietal and allocentric hippocampal systems undoubtedly
work together, not least because all sensory information enters the brain
in an egocentric form and the read-out of an allocentric map must at
some stage be translated into egocentric motor commands such as
turning left or right. Indeed, there is evidence that the firing of neurons in
area 7a of parietal cortex (the part that projects to parahippocampal
cortex) also reflects stimulus locations relative to the monkey’s
environment (Snyder et al. 1998). The complementary roles of these two
brain regions may also be understood in terms of the time scales over
which they support behavior. The hippocampus is involved in long-term
memory, whereas the parietal lobe is involved in perception and action
over short time scales. It makes sense to use an allocentric representation
to store spatial locations over the long term, as the position of the body
will have changed before the information is to be used. However, to act on
a spatial location (e.g., to reach with the hand or look with the eye) an
egocentric representation (e.g., the position relative to the hand or eye) is
more useful (see Burgess et al. 1999).
The rat’s place cell representation of space appears to be derived from an
appreciation of the relative positions of the walls of its enclosure (O’Keefe
and Burgess 1996), presumably represented in the parahippocampal
cortices that provide the input to the hippocampus. Functional
neuroimaging in humans has also implicated parahippocampal cortex in
the perception of space generated by visual stimuli such as enclosing
walls (Epstein and Kanwisher 1998). Lesions to the medial temporal lobe
(particularly those including parahippocampal cortex) can impair the
recognition of landmarks or scenes (see Farrell 1996, Aguirre and
D’Esposito 1999). Finally, we should note that navigation to a place can be
contrasted with simply learning a sequence of body turns. In rats, learning
to go to a place is often learned rather quickly, such that the rat will
approach the learned place whether this requires a left or right body turn.
However, after several trials of making a particular turn, the turn response
will tend to predominate: the rat will make that turn whichever place it
leads to. Interestingly, while hippocampal lesions tend to impair place
navigation, lesions to the caudate nucleus (part of the basal ganglia)
prevent development of a predominant body-turn response (Packard and
McGaugh 1996), consistent with a role in storing well learned routes.