62 | SCIENTIFIC AMERICAN | SPECIAL EDITION | SPRING 2023

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 THE
BRAIN’S
SOCIAL
ROAD
MAPS
Neural circuits that track our whereabouts
in space and time may also play vital roles
in determining how we relate to other people
By Matthew Schafer and Daniela Schiller
Illustration by Richard Borge

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© 2023 Scientific American

MAKING SENSE O
 F THE WORLD

W e are of ten told that there are no shortcuts in life.

But the brain—even the brain of a rat—is wired in a way that
completely ignores this kind of advice. The organ, in fact, epit-
omizes a short­cut-finding machine.
The first indication that the brain has a knack for finding
alternative routes was described in 1948 by Edward Tolman
of the University of California, Berkeley. Tolman performed
a curious experiment in which a hungry rat ran across an un-
painted circular table into a dark, narrow corridor. The rat
turned left, then right, and then took another right and scurried
to the far end of a well-lit narrow strip, where, finally, a cup of
food awaited it. There were no choices to be made. The rat had
to follow the one available winding path, and so it did, time and
time again, for four days.
must have remembered the location of the food, inferred the an-
gle to it and chosen the route most likely to bring it to its goal.
Quite simply, it must have built a model of the environment.
Such model building or mapmaking extends to more than
physical space. Mental maps may exist at the core of many of our
most “human” capacities, including memory, imagination, infer-
ences, abstract reasoning and even the dynamics of social inter-
On the fifth day, as the rat once again ran straight across the actions. Researchers have begun to explore whether mental maps
table into the corridor, it hit a wall—the path was blocked. The an- document how close or distant one individual is to another and
imal went back to the table and started looking for alternatives. where that individual resides in a group’s social hierarchy. How
Overnight, the circular table had turned into a sunburst arena. In- does the brain, in fact, create the maps that allow us to make our
stead of one track, there were now 18 radial paths to explore, all way about the world?
branching off from the sides of the table. After venturing out a few
inches on a few different paths, the rat finally chose to run all the A SPATIAL MAP
way down path number six, the one leading directly to the food. The first hints of a neural basis for mental maps came in the
Taking the path straight to the food cup without prior expe- 1970s. While studying a brain region called the hippo­campus in
rience may seem trivial, but from the perspective of behavioral rodents, John O’Keefe of University College London, along with
psychologists at the time, the rat’s navigational accomplishment his student Jonathan Dostrovsky, discovered a particular class of
was a remarkable feat. The main school of animal learning in neurons that becomes active when mice occupy specific locations
that era believed that maze behavior in a rat is a matter of sim- in their environment. Some of these neurons fired when the ani-
ple stimulus-response associations. When stimuli in the environ- mal was in one location, and others switched on when it moved
ment reliably produce a successful response, neural connections to the next spot on the path along which it traveled, as if the cells
that represent this association get strengthened. were specialized to track where t he animal was in space. By link-
In this view, the brain operates like a telephone switchboard ing sequences of these “place cells” together, researchers were able
that maintains only reliable connections between incoming calls to reconstruct an animal’s navigational trajectory. Work over the
from our sense organs and outgoing messages to the muscles. But intervening decades confirmed the existence of place cells in oth-
the behavioral switchboard was unable to explain the ability to er animals, including humans, and clarified many of their prop-
correctly choose a shortcut right off the bat without having first erties. Along the way, a host of cell types surfaced, each uniquely
experienced that specific path. Shortcuts and many other intrigu- contributing to the brain’s encoding of spatial representations.
ing observations along these lines lent support to a rival school of In the nearby entorhinal cortex, a region connected to the
thought promulgated by theorists who believe that in the course hippo­campus, a research team led by Edvard Moser and May-
of learning, a map gets established in a rat’s brain. Tolman—a pro- Britt Moser, former postdoctoral visiting fellows in O’Keefe’s lab-
ponent of that school—coined the term: the cognitive map. oratory, discovered neurons highly similar to place cells. These
According to Tolman, the brain does more than just learn the cells also fired when an animal was in specific locations. But un-
direct associations among stimuli. Indeed, such associations are like place cells, each of these newly discovered cells spiked in mul-
often brittle, rendered outdated by changes in the environment. tiple, regular locations. When mapped onto the animal’s position,
As psychologists have learned in the decades since Tolman’s work, the activity patterns of these “grid cells” resembled highly regu-
the brain also builds, stores and uses mental maps. These models lar, equilateral triangles. Like a spatial metric, these cells fired
of the world enable us to navigate our surroundings, despite com- when an animal passed over the vertices of the triangles. The dis-
plex, changing environments—affording the flexibility to use short- covery of these cell types sparked excitement because of the
cuts or detours as needed. The hungry rat in Tolman’s experiment emerging picture of how the brain controls navigation. Place cells

64 | SCIENTIFIC AMERICAN | SPECIAL EDITION | SPRING 2023

© 2023 Scientific American

 Giving Way to the Abstract
Maps simplify the world b  y reducing an overwhelming amount of sensory and cognitive data into a format that can be used for
navigating physical space, pointing to shortcuts and detours to reach a destination faster. The organization of such maps—built on
the activity of cells dedicated to tracking both space and time—scales in the abstraction of what they represent: from the recognition
of another individual along the way to even a complex space that denotes social power and closeness to others.

Increasing Abstraction
Physical space Social information in physical space Social space
More

Self
Power
y y

Less
x x Close Affiliation Distant

and grid cells could provide a means to locate oneself in space Shaping of plans also occurs during sleep. Sequences of place
and determine distance and direction. These navigational tools cell activity can be reactivated during sleep to replay the past or
are crucial for building mental maps. (O’Keefe and the Mosers simulate the future. Without the ability to simulate new behav-
received the 2014 Nobel Prize in Medicine or Physiology for their iors, we would have to explore a multitude of real-world options
work on place and grid cells.) before deciding on what action to take. We would be constant
A wide variety of information is useful for creating such a map, empiricists, able to act only on direct observations. Instead off-
and the hippo­campus-entorhinal system encodes much of it. Dis- line simulations give us the ability to envision possibilities with-
covering the location of a physical goal is one example: as an ani- out directly experiencing them.
mal navigates toward an objective, some hippocampal neurons fire
depending on the direction and distance to reach it. The cells in- MENTAL TIME TRAVEL
crease their firing rate as the animal approaches the goal. Time and space are inextricably linked. It is difficult to talk about
Other cells also enter the picture. A dedicated population of “re- time without borrowing a spatial metaphor: time “passes” as we
ward” cells encodes reward locations across different environments, “move” through it. We look “forward” to the future and “back” on
providing a signal to guide an animal’s navigation (think of an “X” our past. The same hippocampal-entorhinal system tracks move­
marking the spot of treasure on a pirate’s map). Other cells track ment through time. Work done largely in the lab of the late How-
speed and direction and in doing so act like internal speedometers ard Eichenbaum of Boston University re­­vealed neurons in the
 ol. 100; October 24, 2018

and compasses that compute an animal’s progress as it travels hippocampal-entorhinal system that encode the time course of
through the environment. Specific cells that signal the locations of an animal’s experience. Time cells fire at successive moments
landmarks in the surroundings serve as references to correct er- but do not track time in a simple clocklike fashion. Instead they
rors in the animal’s trajectory. A map must also have edges: cells mark temporal context—stretching or shrinking their firing du-
that fire more as the animal approaches the map’s perimeter. rations if the length of a task changes, for example. Some time
Source: “Navigating Social Space,” by Matthew Schafer and Daniela Schiller, in N euron, V

For humans, the importance of such an abundance of cell types cells encode space as well. In the brain, in fact, physical and tem-
seems obvious: the brain is responsible for knowing the location of poral space may be bound together.
home and work, walls and dead ends, a favorite shop or the corner The discovery of the crucial importance of these brain areas
store. It is still a mystery as to how all of this information is drawn in space and time was not totally surprising. Psychologists had
together into a coherent map, but these cells appear to provide the long suspected it to be the case. In 1953 Henry Molaison under-
parts list for the elements of neural mapmaking. went bilateral hippocampal resection surgery to reduce ex­­treme,
This hippo­campal-entorhinal system is more than a mapmak- life-disrupting epileptic seizures. The surgery was successful at
er, though, and the maps are more than a way to locate oneself in quelling the seizures. But Molaison—known for decades only as
space. These maps also are used for active planning. When a rat H.M.—became one of the most renowned cases in the history of
comes to a junction in a familiar maze, it will pause while place the brain sciences.
cell firing sequences that relate to the different options are acti- Molaison could remember most experiences from before his
vated, as if the animal is contemplating the choices. surgery—people he knew and recollections from culture and
Humans engage similar processes. Research in participants politics. But his ability to form such explicit memories postsur-
navigating virtual environments while their brains were scanned gery was practically nonexistent. Even so, certain types of learn-
with functional magnetic resonance imaging shows that the ing and memory remained untouched: he could still learn some
hippo­­campus becomes active in ways consistent with spatial new skills with enough practice. But his recollections of new
planning, such as considering and planning routes. people, facts and events were immediately lost.

Graphics by Jen Christiansen SCIENTIFICAMERICAN.COM | 65

© 2023 Scientific American

 Cognitive Cartography
Locations that Is Physical a
 nd S
 ocial
prompt place
The brain forms the idea of friend or foe by stitching together
Hippocampus cell firing
(home of diverse social characteristics from memories that track one’s
place cells) whereabouts. The recollections, research suggests, can then
Locations that be used to place an individual within a social hierarchy that
Entorhinal cortex prompt grid elucidates, say, where one stands in relation to others.
(home of grid cells) cell firing

PLACE AND GRID CELLS MAKING THE LEAP TO SOCIAL MAPS

Place cells pinpoint the animal’s whereabouts, each cell firing when Go right at the corner and continue to your destination. Building
a particular spot on a mental map is reached. A grid cell activates a map of physical surroundings is the work of place and grid
when an animal passes over the vertices of triangles superimposed cells. But the brain may also use these cells for constructing maps
on a mental map. The triangles’ pattern of activation helps the animal for social milieus: locating an acquaintance who grows closer but
compute the direction and distance traveled along a route. loses power in a relationship.

Physical Navigation Social Navigation

y More

Self
Power

Less
x Close Distant
Affiliation

From observing Molaison, neuroscientists discerned that the making. Mapping allows relations to be inferred, even when they
hippo­campus was essential in forming the episodic memories have not been experienced. It also allows for mental shortcuts

 ol. 100; October 24, 2018

that record facts and events. Research on the role of the hippo­ that go beyond the purview of the spatial and temporal domains.

Sources: “Scientific Background: The Brain’s Navigational Place and Grid Cell System,” by Ole Kiehn and Hans Forssberg, with illustrations
campus in episodic memory exploded, largely in parallel to stud- In fact, reasoning using abstract concepts may depend on some
ies on its maplike functions. of these same neural foundations.
The discoveries about the roles of the hippo­campus and en- In one example of this line of work, researchers Alexandra
torhinal cortex in spatial navigation and episodic memory were Constantinescu, Jill O’Reilly and Timothy Behrens, all then at

by Mattias Karlen. Nobelprize.org; “Navigating Social Space,” by Matthew Schafer and Daniela Schiller, in N euron, V
significant for at least a couple of reasons. The work in spatial the University of Oxford, asked participants to learn associations
navigation in rodents marked the first time that a higher-order of different symbols with images of “stick” birds with various
cognitive function—something beyond basic sensory processes— neck and leg lengths. A bird with a long neck but short legs, for
mapped onto clear neural correlates. H.M. showed us that there example, might be linked with the image of a bell, whereas a bird
were multiple types of memory supported by at least partially with a short neck and long legs might be connected to a teddy
different neural systems, with the hippo­campus playing a cen- bear. These linkages created a two-dimensional association space.
tral role in the formation and storage of new episodic memories. Despite neuroimaging being too crude to detect actual grid cells
These discoveries hinted that mechanisms of spatial and tempo- in the human brain, imaging conducted during the learned-
ral navigation might underlie episodic memory. This synthesis association testing nonetheless revealed a gridlike pattern of ac-
is perhaps best ex­­plained by the theoretical construct proposed tivation within the entorhinal cortex.
decades earlier by Tolman; both episodic memory and spatial This finding builds on earlier work by Christian Doeller of the
navigation might reflect the brain’s formation and use of cogni- Max Planck Institute for Human Cognitive and Brain Science in
tive maps. Leipzig, Germany, and Neil Burgess of University College Lon-
Maps are not accurate portraits of the world in all of its com- don that first showed an entorhinal gridlike representation in
plexity. Rather they are representations of relations—distances humans navigating a virtual maze. For both physical and abstract
and directions between locations and what exists where. Maps relations, the gridlike organization is highly efficient. It makes
reduce a dizzying amount of real-world information into a sim- the linkages of places or concepts more predictable, enhancing
ple, easily readable format that is useful for effective, flexible nav- how quickly inferences can be made about these relations. As in
igation. The cell types mentioned earlier (place cells, grid cells physical space, this organization of information allows for the
and border cells, among others) may piece together such related inference of shortcuts—relations between ideas or perhaps anal-
elements into a mental map, which other brain regions can then ogies, stereotypes and even some aspects of creativity itself could
read out to guide “navigation,” amounting to adaptive decision- depend on such inferences.

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 MAKING SENSE O
 F THE WORLD

PEOPLE MAPS dimensions of power and affiliation. In each interaction, we drew

The progression f rom the physical to the abstract carries over a line or vector from the participant to the character. In this way,
into the way the brain represents social relationships. Various we charted the evolving relations as trajectories through social
bits of knowledge about another person are distilled into the space and computed information about the angles and lengths
concept of that individual. When we see a photograph of some- of the social vectors.
one or hear or see that person’s name, the same hippocampal We searched for neural signals that tracked this information
cells will fire, regardless of the sensory details of the stimulus by correlating a participant’s brain activity with the angle and
(for example, the famous “Jennifer Aniston neuron” de­­scribed length of the vectors for each decision. Activity in the hippo­
by Itzhak Fried of the University of California, Los Angeles, and campus tracked the angle of the characters to the participant.
his colleagues). These hippo­campal cells are responsible for rep- The degree to which hippocampal activity captured these social
resenting concepts of specific individuals. coordinates also reflected the participants’ self-reported social
Other hippocampal cells track the physical locations of oth- skills. These findings suggest that the hippo­campus monitors so-
ers and are called social place cells. In an experiment by David cial dynamics as it does physical locations by encoding relations
Omer of the Hebrew University of Jerusalem, Nachum Ulanovsky be­­tween points in multidimensional space. Indeed, it may be
of the Weizmann Institute of Science in Rehovot, Israel, and their that along any arbitrary dimension in which we can order
colleagues, bats observed other bats navigating a simple maze information, whether physical or abstract, the hippo­campus-
to reach a reward. The task of an observer bat was to simply entorhinal system plays a part.
watch and learn from a navigating bat, enabling it to subsequent- Many questions about the brain’s social maps still remain un­­
ly navigate the same route to get the same reward. When the ob- answered. How does this system interact with other brain re-
server bat watched, hippocampal cells fired corresponding to gions? For example, in our role-playing study, we found that the
the location of the other bat. posterior cingulate cortex, a region also involved in represent-
Neural circuitry within specific subregions of the hippo­ ing spatial information, tracked the length of social vectors—
campus (in particular, areas called CA1 and CA2) contributes to functioning in effect as a measuring stick of “social distance.”
such social memories. Artificial stimulation or inactivation of Further, a gridlike signal was found in brain regions that are in-
these hippocampal areas enhances or diminishes an animal’s terconnected with and tend to co-activate with the hippo­campal-
ability to recognize other animals. In humans, hippocampal entorhinal system, suggesting they form a network of brain re-
­injury often spares memory for specific, individual faces, but gions with common functional properties.
the relation between this cardinal identifier of another person As research accumulates, questions of clinical importance
and that individual’s behavior may be lost. That observation sug- arise as well. Can flawed mapping processes explain psychiatric
gests that the hippo­campus does not simply record a face or dysfunction? Another possibility is that insights garnered from
some other personal detail but rather ties together diverse so- this brain architecture could inform artificial-intelligence devel-
cial characteristics. opment. Well-organized internal models of the world might be
Hippocampal activity also tracks social hierarchies: the de­­ key to building more intelligent machines.
mands of a boss and a co-worker, for instance, may be valued dif- That the same mapping system may underlie navigation
ferently and confer different social standings. Common meta- through space and time, reasoning, memory, imagination and
phors illustrate the spatial dimensions of a hierarchy: a person even social dynamics suggests that our ability to construct mod-
may try to gain status to “climb the social ladder” or “look down” els of the world might be what makes us such adaptive learners.
at someone below them. Other factors are also critical. Biologi- The world is full of both physical and abstract relations. Road
cal relatedness, common group goals, the remembered history maps of city streets and mental maps of interrelated concepts
of favors and slights—all determine social proximity or distance. help us make sense of the world by extracting, organizing and
Human relationships can be conceived of as geometric coordi- storing related information. A new coffee shop on a familiar
nates in social space that are defined by the dimensions of hier- street can be easily placed within an existing spatial map. Fresh
archy and affiliation. concepts can be related to older ideas. And a new acquaintance
Work in our lab has explored these ideas in recent years. Our can reshape our social space.
results suggest that, as with other spaces, the hippo­campus or- Maps let us simulate possibilities and make predictions, all
ganizes social information into a map­like format. To test this hy- within the safety of our own heads. The mental shortcuts we
pothesis, we put individuals in a choose-your-own-adventure can so readily conjure up might have their basis in the same sys-
game in which they interacted with cartoon characters and made tem that allows us to figure out a detour around a traffic jam.
decisions while their brains were scanned. We have just begun to discover the varied properties and capac-
In the game, players had just moved to a new town and need- ities of this system. Mental maps do more than help us find
ed to interact with the fictional characters to secure a job and shortcuts through physical space—they enable us to navigate
a place to stay. Participants made decisions on how to deal with life itself.
a given character. Players could request that others perform fa-
vors to demonstrate their power, or they could submit to de- Matthew Schafer is pursuing a doctorate in neuroscience at the Icahn School of Medicine
mands made on them. In a subsequent interaction, they could at Mount Sinai, focusing on the neural mechanisms of social cognition in the human brain.
decide whether to make a gesture of attachment—giving a hug
or remaining at a distance. Daniela Schilleris a professor of both neuroscience and psychiatry at the Icahn School
Using these decisions, we plotted each character at certain of Medicine at Mount Sinai. She researches the neural mechanisms underlying emotional
coordinates on a map representing their movement along the control needed to adapt to constantly changing environments.

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