Spatial Cognition
Navigation: Finding the way to a goal
Discriminate different headings (need a sense of direction)
• External directional reference: sun, magnetic field, landmarks
• Internal directional reference: vestibular/inertial cues
Determine the correct heading (need a sense of position)
• Path integration
• Knowledge of familiar landmarks in home range
• Geographical positioning system (e.g., position relative to large-scale
coordinate system defined by global geophysical features)--migratory
birds, whales, turtles
Spatial Cognition and Navigation
Navigational Processes That Use Internally
Represented Spatial Knowledge
• Path integration
• Sun compass
• Landmarks: cognitive maps
Path integration: a sense of position
• Ants travel straight path home after
circuitous outward path
• We can exclude use of odor trails,
visual beacons, or memory of
outward path
• Instead, ants compute direct
homeward path based on
measurements of directions and
distances traveled on outward path
(Wehner et al.)
Food
Homeward
path
Nest
More on PI
Animals compute straight path home after circuitous outward path
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Food
Search
path
Desert ants
(Cataglyphis sp.)
Homeward
path
Nest
QuickTime™ and a TIFF (LZW) decompressor are needed to see this picture.
http://axon.bhs.mq.edu.au/PSY236/space/20path_int_error_2.jpg
www.kyb.tuebingen.mpg.de/bu/poster/ 2000/b_riecke_arvo2000.pdf
Sun Compass and Memory in Bees
The basic task
H
16:00
16
12
Noon
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F
http://www.scottcamazine.com/photos/
BeeBehavior/images/06waggleDance_jpg.jpg
• Bees encode (allocentric?) flight direction in dances
• As sun moves, dances change
• Dances change even when bees can’t see sun (thus
compensate by memory)
• Reference for memory: landmarks (Dyer & Gould
1981; Dyer &Dickinson 1996)
Food
20°
40°
75°
(Up)
20°
40°
75°
Celestial compasses: birds and bees

Inputs
Outputs
Sola r Azim uth
Time of Day 
Sun
Landmarks
Time
0
45
90
135
180
225
270
315
360
 = f ()
21
18
15
12
9
6
3
(Renner 1959; Dyer 1987)
Relative
 = f ()
Azimuth 
(Beier and Lindauer 1970)
Pigeons
Budzynski, Dyer & Bingman 2000
Landmark Panoram a
Food
20°
40°
75°
(Up)
Pattern of solar movement
• Non-linear over day
• Varies with season and latitude
• Animals learn current, local pattern of solar
movement
• Learning: not just a list of time-linked solar
positions, but a function that can be used to find
unknown positions of the sun.
20°
40°
75°
Birds naïve about the morning
sun can nevertheless use it
correctly based to find compass
directions learned in afternoon
Solar
positions
seen
Food
14:00
Training
12:00
To get food in box, birds must
chose correct angle relative to
sun, compensated for solar
movement
10:00
(Test)
Landmarks: Cognitive Maps
From: Tolman, EC 1948 Cognitive maps of rats and men. Psychol Rev 40: 40-60.
What does it mean to have a cognitive map?
One operational definition: a representation of
spatial relationships that enables computation of
novel shortcuts between known locations (O'Keefe
& Nadel 1978. The hippocampus as a cognitive
map)
Alternative hypotheses:
• Route memory (A--> B already familiar)
• Recognize familiar landmarks associated with
goal, even if from novel vantage point
Task: get from A to B, having
experienced routes to A and B
separately
Learning local landmarks
Insects can pinpoint locations they need
to find again by learning arrangement of
surrounding landmarks: HOW?
Niko Tinbergen (1938)
Learning local landmarks
Bees match visual image learned on
previous trips
Find best match given all available
information
Search
distributions
But bees have some
flexibility in approach
path. They don’t follow
stereotyped route, which
Gallistel takes as evidence
of generalized “map”
Landmark
Enlarge single landmark; to match Enlarge three landmarks; best
view, bees have to move back
match is in the same place
(Bartlett & Dyer in prep)
Algorithm: snapshot model
Insect records image of
landmarks seen at goal
(after Cartwright and Collett 1983)
Vardy www.scs.carleton.ca/~avardy/ misc/engrSeminar/engrSeminar.pd
Then finds goal on later
approach flights by
moving to reduce
mismatch between
current and remembered
images
Is this evidence for a map?
Robotic simulations
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Source: Möller, R., Universität Bielefeld
http://www.ti.uni-bielefeld.de/html/people/moeller/analog.html
But is this really what bees do?
Or this?
Bartlett, Mack & Dyer (in prep)
A simpler model?
• Bees encode angles (and distances) of
landmarks; may be encoded in egocentric, not
allocentric, reference frame
• Weak evidence for a highly flexible
computational strategy for using landmarks to
fly to goal
• Nevertheless, bees do behave as if they can
recognize familiar landmarks from novel
vantage points
• Also, bees can use familiar landmarks
encountered in unexpected context
What does it mean to have a cognitive map?
One operational definition: a representation of
spatial relationships that enables computation of
novel shortcuts between known locations (O'Keefe
& Nadel 1978. The hippocampus as a cognitive
map)
Alternative hypotheses:
• Route memory (A--> B already familiar)
• Recognize familiar landmarks associated with
goal, even if from novel vantage point
Task: get from A to B, having
experienced routes to A and B
separately
Do insects have cognitive map or something else?
N
50 m
A
C
H
F
265
262
HIVE
259
F
C
(H)
B
From Dyer 1991
Varieties of cognitive maps? (Gallistel 1990)
Broader Definition (Gallistel 1990): ‘A cognitive map is a record in the
central nervous system of macroscopic geometric relations among surfaces
in the environment used to plan movements through the environment. A
central question is what type of geometric relations a map encodes’.
Specific issues:
• Spatial scale (local vs. home-range)
• Geometric content (metric, topological)
• Reference frame (egocentric/view-dependent vs. allocentric/viewindependent)
Evidence:
• People: short cuts in cities and VR (errors); mixed evidence contents of
underlying map
• Rodents: most studies on local scale; mixed evidence on contents
• Insects: on local and home-range scale--metric, egocentric
Varieties of cognitive maps?
Type 1
Local Image (Snapshot)
Type 2
Type 3
Route Maps
Global (Metric) Map
F1
F1
N
F2
F2
Experienced
Computed
Most rodent research
Humans in rooms
•
•
•
Insects (digger wasps, bees)
Humans in corridors, cities
`
Humans in cities
Computational models of cognitive maps: need to specify geometric contents (angles,
distances, routes, nodes), reference frames, and operations performed on stored
information?
Humans, but not insects, form Type 3 maps, but insects can flexibly use snapshots and route
maps
Big question is whether map-learning is viewpoint-dependent or viewpoint-independent
Toward the implementational level
Is the hippocampus the locus of the cognitive map of mammals?
Rat brain
Some evidence:
• Input from integrative sensory areas
• Output to neocortex
• Lesion studies suggest role in memory
CA1
Section through hippocampus
http://www.sunysb.edu/biochem/BIOCHEM/facultypages/trimmer/gallery.html
CA3
Hippocampus and spatial cognition
1. Lesion experiments (rats & birds): selective effect on
spatial memory
2. Comparisons of hippocampus size: correlation
between HC size and reliance on spatial memory
3. Functional neuroimaging (humans)
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Maguire, E.A. 1998
Science 280: 921-924
4. Place cells
Normal (top) and HClesioned brains
http://www.psychol.ucl.ac.uk/kate.jeffery/C5
67/Lecture2_Cognitive_mapping/sld001.htm
Place Cells
Firing field of single cell (gray)
and activity on two runs
through the arena. Cell's
activity is independent of
heading. Muller, R. (1996) A
quarter of a century of place
cells. Neuron 17: 813-822
Rat with recording apparatus
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Rat in arena
Ensemble of hippocampal cells "maps"
space. This shows the place fields of five
simultaneously recorded units on an
elevated triangular maze. The light gray
trace represents the rat’s path as it traversed
the maze. The dots in each color indicate
the locations in which each of the five cells
fired. (Best et al. 2000, Ann Rev.
Neuroscience 24:459-86; data from C
Barnes & B McNaughton.)
Place Cells (cont'd)
Rat put in dark and then rotated slowly relative
to featureless arena; causes shift in place field
Conspicuous landmark is rotated, causing shift
in place field
Place fields are referenced to
internal and external coordinates
Remapping of place fields of when animal goes
into different environments; one cell's place
fields are shown
Some properties of hippocampal place cells that
imply role in spatial cognition
• Highly stable firing fields in constant environment
• Can be established in total darkness (referenced to vestibular
cues)
• Can be linked to visual cues, and then track visual cues
• Individual cells can have different place fields in different
environments
• Ensemble of cells encodes map of familiar environment
• BUT: there is no obvious way in which this system
corresponds to computational models of cognitive map based
on behavioral evidence. For example…..
Problems with the "hippocampus-as-cognitivemap" hypothesis
• May not generalize to humans, because hippocampus is
known to play role in non-spatial episodic memory in
humans (but see O’Keefe)
• Place cell ensemble in hippocampus is not enough to account
for spatial behavior….encodes current location, but not goals,
for example (Andre Fenton)
• Even in animals, hippocampus is involved in non-spatial
tasks (e.g., transitive inference)
• Place cells seem to encode something more than just "place"
Transitive inference: non-spatial function of
hippocampus in rats
• Rat chooses one odor over another
IF: A > B > C > D > E, then….
A > C (or D or E); B > D (or E)
• Rat can learn series of pairwise
discriminations
• Rat can perform transitive inference,
but not if hippocampus is damaged
Normal rats
HC-damaged
Eichenbaum, H 1999 Behav Brain Res 103: 123-133
Place cells encode more than just place:
role for hippocampus in episodic memory?
Rats are trained on
alternating T-maze
Wood ER, Dudchenko PA, Robitsek RJ, Eichenbaum H: Hippocampal neurons encode
information about different types of memory episodes occurring in the same location.
Neuron 2000; 27: 623-633
Context-specific activity of place cells
A cell that fires in a
particular place, but
much more rapidly when
a right turn is coming up
A cell that fires in
different locations when
a right turn is coming up
than when a left turn is
coming up
Wood ER, Dudchenko PA, Robitsek RJ, Eichenbaum H: Hippocampal neurons encode
information about different types of memory episodes occurring in the same location.
Neuron 2000; 27: 623-633
Episodic Memory in Animals?
Episodic Memory in Birds:
where did I put that worm and when did I put it there?
Birds are allowed to hide food
(waxworms and nuts) in two caching
bouts. In recovering it they can choose
based on what they cached and when,
and their knowledge of what happens
to each food type over time.
http://freespace.virgin.net/cliff.buckt
on/Birding/California/Calif17.jpg
Clayton & Dickinson 1998
Worms Degrade
Worms Stay Tasty
Worms Disappear
Interpretations for role of hippocampus in spatial
cognition and memory generally
• Spatial and non-spatial episodic memories involve different processes;
hippocampus does them both, and has evolved to become more
specialized for episodic memory in primates compared with rodents
(Jacobs & Schwenk)
• All experience has a spatial component, and hippocampus participates
in formation/use of episodic memory because of its role in processing
spatial information processing (O'Keefe)
• Spatial cognition involves processes that are also required in encoding
certain other kinds of relations among stimuli; hippocampus plays a
more general role that leads it to participate in spatial as well as certain
non-spatial tasks (Eichenbaum)
Where does this leave the search for neural
implementation of cognitive map?
Hypotheses
• Hippocampus is the cognitive map (O’Keefe)
• Cognitive map (sensu Tolman and O'Keefe & Nadel) is
elsewhere, but uses output from hippocampus
• Cognitive map is indeed an important function of
hippocampus, but computations that hippocampus carries out
are very different from those developed on the basis of
behavioral observations; these computations support functions
other than spatial encoding (Eichenbaum and colleagues)