Neuron
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The Hippocampus
as a Cognitive Map . of Social Space
Howard Eichenbaum1,*
1Center for Memory and Brain, Boston University, Boston, MA 02215, USA
*Correspondence: hbe@bu.edu
http://dx.doi.org/10.1016/j.neuron.2015.06.013
The traditional view of the hippocampus is that it creates a cognitive map to navigate physical space. Here, in
this issue of Neuron, Tavares et al. (2015) show that the human hippocampus maps dimensions of social
space, indicating a function in the service of navigating everyday life.
Edward C. Tolman (Tolman, 1948) devel-
oped the notion of cognitive maps as
a heuristic for understanding the complex
cognitive mechanisms that guide
behavior. His theory of purposeful
behavior was aimed to contrast with
the contemporaneous, widely accepted
view that behavior is guided by stimulus-
response connections, and his experi-
ments identified specific abilities that re-
flected cognition outside the scope of
behavior that could be supported by stim-
ulus-response learning. Tolman’s experi-
ments focused on rats solving maze prob-
lems, but he did not interpret his findings
narrowly as a description of navigational
computations. Rather, he employed
spatial learning to model aspects of
goal-oriented decision-making, and he
viewed a cognitive map as an organiza-
tion of cognitive operations. Tolman
emphasized that cognitive maps provide
insights into human cognition broadly,
including human social behavior. In this
issue of Neuron, Tavares et al. (2015)
realize Tolman’s broader view of cognitive
maps by their characterization of a cogni-
tive map of social organization in humans
supported by the hippocampus.
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A connection between the hippocam-
pus and cognitive maps began with a
landmark book by O’Keefe and Nadel
(1978), who proposed that the hippocam-
pus provided the neural basis of cognitive
mapping. The book, and decades of ex-
periments and theoretical work that fol-
lowed, departed from Tolman’s map of
cognition to instead focus on cognitive
maps as psychological and neural repre-
sentations of physical space, and on
mechanisms within the hippocampus
and associated brain areas that create
geographical maps and perform navi-
gational computations (Hartley et al.,
2014). Originally, O’Keefe and Nadel
(1978) extended their model to the repre-
sentation of items and events in spatial-
temporal context as an evolutionary
advance of the human hippocampus to
support its function in mapping mem-
ories, particularly as it might represent
the deep structure of language. This
extension of their theory clearly went
beyond physical space (although only
for humans) and, in doing so, acknowl-
edged that information processing by
the hippocampus can in principle be
applied outside the domain of physical
Neuron 87, July 1, 2015 ª2015 Elsevier Inc. 9
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space. To what non-spatial domains
does it apply?
Tavares et al. (2015) here reveal that
cognitive maps in the hippocampus
extend to social space. Social space is
an excellent candidate for hippocampal
representation, because it is a domain
that, like physical space, is characterized
by a combination of continuous dimen-
sions. So, just as planar geography is
characterized by two spatial dimensions,
social space has been characterized by
multiple social dimensions, including of
particular relevance here, the dimensions
of power and affiliation. Importantly, just
as the dimensions of geographic space
are defined in terms of continuous metrics
of physical distance, power and affiliation
are defined in terms of continuous metrics
of social distance.
To test the idea that social relations
are mapped within the hippocampus, Ta-
vares et al. (2015) designed a role-playing
game in which participants imagined
they had moved to a new town and their
goal was to find a job and place to live.
To accomplish this, the participants
conversed with local people in the search
for a job or home through different
 Neuron

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responses in which they could hippocampal volume gray

comply with a character’s de- matter is increased during
mand or make demands medical training, although
(increasing or decreasing the Woollett et al., (2008) reported
power of the character) and that medical doctors do not
engage or not engage in per- have larger hippocampi,
sonal conversation and phys- whereas London taxi drivers
ical interaction (increasing or do. Most intriguing is a report
decreasing affiliation with the that professional piano tuners
character), and the accumula- who spend thousands of hours
tion of successive interac- navigating acoustic scenes
tions took the narrative in have larger hippocampal gray
different directions. The out- matter volumes associated
comes of these social interac- with the history of training
tions positioned each char- Figure 1. Cognitive Maps of Physical and Social Space experience (Teki et al., 2012).
The map of physical space is Cartesian and allocentric, referenced to
acter along axes of power the geometry of the environment and not to the individual observer. By
Studies on hippocampal vol-
and affiliation measured as contrast, the map of social space is polar and egocentric, referenced to ume do not inform us about
relational distance to the the observer. the nature of hippocampal
participant, and the combina- representation but the paral-
tion of relational distances lels between characterizations
along power and affiliation axes consti- to consider, however, that hippocampal of systematic mapping of multiple dimen-
tuted a vector describing the participant’s neural activity also maps paths through sions across musical, spatial, temporal,
social relationship to each character in po- space, including distinct representations and now social structure are striking.
lar space (Figure 1). As interactions pro- of paths that overlap in space, which The Tavares et al. (2015) findings also
ceeded in the narrative, the position of therefore measure the animal’s personal parallel evidence that the hippocampus
the character moved along the coordi- experience within an allocentric frame- is essential to relational organization in
nates of these two dimensions. To test work (Wood et al., 2000). Furthermore, animals and humans as they employ a
their hypothesis that the hippocampus hippocampal neural activity also maps systematic mapping of relations between
maps social space, Tavares et al. (2015) the temporal organization of experience, arbitrary stimuli. Early studies showed
used a General Linear Model to regress independent of, or along with, movement that rats can learn a set of overlapping
hippocampal activation measured in an in allocentric space, indicating that one’s stimulus associations (A is associated
fMRI signal to the vector in the established sense of time is another self-generated with B; B is associated with C) that sup-
social space. Their main result was that the dimension mapped by the hippocampus port inference between indirectly related
fMRI signal in the left hippocampus corre- (Eichenbaum, 2014). In addition, recent stimuli (therefore A is associated with C),
lated with the vector angle, indicating that evidence suggests that the spatial map indicating the existence of a cognitive
the hippocampal network identified each is topological, that is, according to adja- map of a simple associative space (Bun-
character’s position in social space as cency of locations rather than reflecting sey and Eichenbaum, 1996). Hippocam-
an interaction of their power and affiliation the geometry of an environment (Daba- pal lesions prevent formation of this
relations. Furthermore, the tracking ghian et al., 2014), suggesting that hippo- mapping in animals and, in humans per-
strength for these dimensions correlated campal mapping is more about spatial forming the same task, the hippocampus
with the social skills of the participant, as associations than Cartesian coordinates. activates as the associative space is
assessed in a combination of question- In sum, the perspective, parameters, formed and employed (Zeithamova
naires on social qualities and personality and metrics of organization in hippocam- et al., 2012). Similar reports have shown
traits. pal networks seem to reflect the differing that the hippocampus is essential to the
Notably, the hippocampus was not the dimensions of experience across do- mapping hierarchical relations among
only brain area that mapped social mains of mapping. If there is a universal stimuli (A > B > C > D > E) in animals (Du-
space, just as it is not the only brain format for relational representation in sek and Eichenbaum, 1997) and that the
area that represents physical space. the hippocampus, it remains to be hippocampus is activated in the same
Thus, the hippocampus is best viewed discovered. task in humans (Heckers et al., 2004).
as a ‘‘hub’’ that coordinates widespread Does the hippocampus map all manner These observations strongly complement
cortical areas that map different forms of spaces? The findings of Tavares et al. the findings of Tavares et al. (2015) on so-
of space. Also, it is notable that the map- (2015) parallel reports that the hippocam- cial space, and further extend the range of
ping of egocentrically anchored social pal volume is correlated with exercise in domains in which the hippocampus sup-
space in the Tavares et al. (2015) study mapping and navigating auditory space ports maps of cognition.
differs from the emphasis on mapping al- and medical school training. Studies Eichenbaum and Cohen (2014) sug-
locentric physical space in studies on have linked musical expertise to hippo- gested that the hippocampus supports
hippocampal neuronal activity in animals campal volume (Oechslin et al., 2013). our ability to ‘‘navigate life,’’ by creating
(O’Keefe and Nadel, 1978). It is important Also, Draganski et al. (2006) reported that a network of memories that permit us

10 Neuron 87, July 1, 2015 ª2015 Elsevier Inc.

Neuron
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to traverse new routes through an ab-
stract memory space and solve new
problems in many domains of everyday
life. We suggested that memories could
be related by space and time, as well
as by other meaningful dimensions of
experience. Consistent with this view,
Tavares et al. (2015) have revealed a
systematic mapping of social space by
the hippocampus, complementing evi-
dence from other approaches that sup-
port the idea that Tolman’s map of
cognition applies very well to the hippo-
campus across domains of relational
representation.
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