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REVIEW
Spatial reorientation in large and small enclosures:
comparative and developmental perspectives
Cinzia Chiandetti Æ Giorgio Vallortigara
Received: 18 April 2007 / Revised: 12 October 2007 / Accepted: 3 January 2008 / Published online: 15 January 2008
Ó Marta Olivetti Belardinelli and Springer-Verlag 2008
Abstract Several vertebrate species, including humans,
following passive spatial disorientation appear to be able to
reorient themselves by making use of the geometric shape of
the environment (i.e., metric properties of surfaces and
directional sense). In some circumstances, reliance on such
purely geometric information can overcome the use of local
featural cues (landmarks). The relative use of geometric and
non-geometric information seems to depend upon, among
other factors, the size of the experimental space. Evidence in
non-human animals and in human infants for primacy in
encoding either geometric or landmark information depend-
ing on the size of the environment is reviewed, together with
possible theoretical accounts of this phenomenon.
Keywords Spatial reorientation Geometry
Modularity Space size Human infants Chick
Pigeon Fish
Introduction
Among the wide range of spatial skills that allow animals
to navigate, the ability to reorient after disorientation in a
familiar environment can not be ignored. Spatial reorien-
tation plays an important role in natural conditions when an
animal has to re-establish the spatial relationship between
itself and its environment in order to find a particular
location (irrespective of whether a human-being is required
to return to a shop or to a restaurant or whether a rat has to
go back to its own hole; the underlying basic mechanisms
seem to be quite comparable (Vallortigara 2008). Reori-
entation is a basic adaptive strategy, strictly necessary to
figure out the exact relationship between an organism’s
current position and an already visited target location when
path integration is ruled out. How do organisms deal with
such spatial re-orientation problems?
Strategies and kinds of information
Two major and complementary coding strategies are in use
when an organism encodes the properties of an external
space: an egocentric- and an allocentric-based strategy.
The former is accountable for tracking the position of the
animal by encoding (and continuously updating) distances
and directions from the starting point during travel: it is
also known as path integration or dead reckoning. The
latter is responsible for mapping out locations by referring
to the characteristics of the surroundings, encoding spatial
relationships among cues, which can provide a stable frame
of reference with their relative positions since they do not
change as the animal moves. Each mechanism bears per se
informative properties and they can co-operate. In addition,
by doing so they may provide redundant information.
However, it may also occur, in some circumstances, that
the two systems are not in agreement, i.e., when a move-
ment not controlled by the animal itself displaces it (as
tumbling down a hill) or when there is a lack of external
C. Chiandetti (&)
Department of Psychology and B.R.A.I.N. Centre
for Neuroscience, University of Trieste,
Via S. Anastasio 12, 34123 Trieste, Italy
e-mail: cchiandetti@univ.trieste.it
URL: http://www.psico.univ.trieste.it/labs/acn-lab/eng_p/
e00_home.html
G. Vallortigara
Center for Mind/Brain Sciences, University of Trento,
Corso Bettini 31, 38068 Rovereto, Italy
e-mail: giorgio.vallortigara@unitn.it
123
Cogn Process (2008) 9:229–238
DOI 10.1007/s10339-008-0202-6
orienting cues (for instance in the dark). In those cases, the
internal system formed by inertial, kinaesthetic and ves-
tibular signals is inevitably disrupted and the objects’
relationships to the animal are no longer represented cor-
rectly but leave the external one still unaffected (since the
position and the orientation of the agent are not relevant for
correctly representing object-to-object relationships). Thus,
the allocentric-based mechanism is used to adjust the self-
based one and aspects of the external world must be used to
re-establish orientation as has been well-documented in a
wide variety of studies addressing this reorientation ability
(see Etienne and Jeffery 2004; Cheng 2005; Cheng and
Newcombe 2005 for exhaustive reviews).
To successfully orient and functionally navigate from
one place to another a detailed consideration of the overall
arrangement of spatial features available in the environment
is needed. Usually, at least in our experience as human-
beings, a place ‘‘can be objectively defined by both its
relationship to the environment and its intrinsic character-
istics (local cues)’’ (Poucet 1993). In the natural
environment, namely the complete space surrounding an
individual (global level), local cues refer to all the distinc-
tive features that can be selectively taken into account, and
defined by relationships that link one single characteristic to
each other but that are still located in the ‘‘working space’’.
Of course, there are no physical boundaries that constrain a
local space with respect to a global one. However, because
of the particular indoor paradigm developed by K. Cheng to
study spatial reorientation abilities in animals (Cheng 1986)
the global space perceived by the animal is defined (and
confined) by the shape of the enclosure: i.e., the layout
composed of axes, surfaces and their relative lengths
together with their incident joint points. In the literature,
this geometric layout is referred to as geometric informa-
tion. Another source of information is supplied by the local
cues—non-geometric information in this paradigm—i.e.,
those salient properties of the surfaces such as their colours
and patterns or those features distinct from the surfaces
themselves such as separate beacons located in specific
positions. In natural environments, these two types of cue
are usually correlated although with varying degree from
one environment to another.
There is, then, another source of information that is
usually present while the animal is processing the sur-
roundings: the internal directional sense of left–right which
may be helpful in associating different cues with respect to
their specific and relative position in egocentric coordinates.
The original paradigm and the first results
A powerful experimental paradigm to study spatial reori-
entation abilities was first developed by K. Cheng in 1986.
In his study, rats (Rattus norvegicus) were located inside a
rectangular enclosure and were trained to find food,
available in one corner, in the presence of several visual
and olfactory cues. The correct location was defined by the
availability of both geometric properties of the surfaces
(i.e., shorter and longer walls together with the ‘‘sense’’ for
left–right discrimination) and non-geometric properties
(visual and olfactory cues of the surfaces of the environ-
ment; a schematic representation of the task is shown in
Fig. 1). When the rats were reintroduced into the apparatus,
after having been passively disoriented outside in order to
Fig. 1 Top schematic representation of the geometrical information,
which is available in a rectangular-shaped environment. The target
(filled dot) stands in the same geometric relations to the shape of the
environment as its rotational equivalent (open dot). Metric informa-
tion (i.e., distinction between a short and a long wall) together with
sense (i.e., distinction between left and right) suffices to distinguish
between locations A–C and locations B–D, but not to distinguish
between A and C (or between B and D). Bottom schematic
representation of the non-geometrical information provided
by discrete panels placed in each corner. The target (filled dot)is
unambiguously defined by the presence of a specific and salient
visual cue
230 Cogn Process (2008) 9:229–238
123
remove inertial information (i.e., to avoid internal sense of
direction and location), they searched at an equal rate in the
target corner and in the corner located at 180° rotation from
the target (the indistinguishable opposite position with the
same geometrical properties with respect to the shape of
the enclosure).
Interestingly, rats did not resort to use of non-geometric
cues to solve the ambiguity of the room’s symmetry and
thus to discriminate between the two geometrically iden-
tical positions (A and C in Fig. 1). A combination of the
two sources of information, geometric and non-geometric,
would have led them to a correct choice but rats were able
to take into account both geometric and non-geometric
properties only after repeated trials (i.e., in a reference
memory task). However, even in reference memory tasks,
when the two kinds of information were placed in conflict
(using the so-called affine transformation, in which a dis-
location of the featural cues occurred so that the previously
reinforced cue was located in a novel, geometrically
incorrect position) rats preferred to choose the corners
located in the geometrically correct locations and with the
wrong features, confirming a primary role of geometric
information in reorientation.
Making use of the same paradigm as described for rats,
Hermer and Spelke in 1994 (and again in 1996) tested 18–
24-month-old children in a rectangular room (4 9 6-ft)
with either all four walls white (geometry-only condition)
or with three white walls and one blue wall (geometry plus
feature condition). In the reference memory task, toddlers
reoriented taking into consideration the geometric proper-
ties of the rectangular room in the white-wall testing
situation, but they continued to confuse systematically the
two geometrically equivalent locations in the blue wall
task, when featural information was available. Thus they
were demonstrated as having primarily used (as did rats)
the geometric shape of the environment to find the target
position. In contrast, human adults behaved in a completely
different manner being able to use geometry alone when in
the white-walls condition and to conjoin geometric and
non-geometric information to disentangle between geo-
metrically equivalent locations when tested in the blue wall
test (Hermer and Spelke 1994).
A modular architecture for spatial reorientation?
This pattern of data was accounted for by Hermer and
Spelke (1994) in terms of a modular architecture of cog-
nitive functions (Fodor 1983, 2001). The authors argued
that toddlers rely on an innate ‘‘geometric module’’, an
encapsulated and task-specific process, devoted to a subset
of information, i.e., purely geometrical information. The
evidence that older children (5–7 years old) appeared to be
able to disentangle the room’s symmetry when in the
presence of a blue wall (Hermer-Vasquez et al. 2001), led
the authors to suggest that the development of spatial
language (with use of ‘‘left’’ and ‘‘right’’ terms associated
with description of visual scenes) would be crucial for the
ability of integrating geometric and featural information
(Spelke 2003; Wang and Spelke 2002).
Further evidence for a role of language came from
studies in which human adults were required to solve the
spatial reorientation task while performing a verbal shad-
owing task so as to prevent use of verbal language.
Hermer-Vazquez et al. (1999) showed that, when tested
while performing a verbal shadowing task, human adults
displayed the same pattern of errors as children in the
presence of a blue wall, mirroring the inability of younger
infants to integrate geometric and non-geometric infor-
mation. When tested while performing a non-verbal
shadowing task (i.e., rhythm clapping), the same subjects
appeared able to conjoin successfully the two sources of
information, thus supporting the hypothesis of a role of
language to overcome the geometric module encapsulation
(but see Ratliff and Newcombe 2005; Hupbach and Nadel
2005; Newcombe 2005 for contrasting evidence).
However, although verbal language could be crucial for
the integration of geometric and non-geometric informa-
tion in humans (Carruthers 2002; Hermer-Vasquez et al.
1999), other species, devoid of linguistic capacities, can
nonetheless conjoin geometric and non-geometric infor-
mation in order to reorient themselves. Evidence has been
collected in this regard from a wide range of species,
including fish [redtail splitfins (Xenotoca eiseni): Sovrano
et al. 2002, 2003; goldfish (Carassius auratus): Vargas
et al. 2004], birds [domestic chicks (Gallus gallus): Val-
lortigara et al. 1990, 2004; pigeons (Columba livia): Kelly
et al. 1998) and mammals (rhesus monkeys (Macaca
mulatta): Gouteux et al. 2001; tamarins (Saguinus Oedi-
pus): Deipolyi et al. 2001].
Does size matter? Reorientation in enclosures
of different sizes
Learmonth et al. (2001) duplicated the original experi-
ments of Hermer and Spelke (1994) but found that young
infants successfully reoriented themselves by joining
together geometrical and featural information. However, an
uncontrolled variable across these two studies was the size
of the experimental room used. Evidence of an influence of
the environment size on navigation and on estimations of
distances had been previously noted in children (see
Herman and Siegel 1978; Siegel et al. 1979). Thus, in a
subsequent study, Learmonth et al. (2002) specifically
addressed this issue. They found that toddlers failed to
Cogn Process (2008) 9:229–238 231
123
consider environmental features as landmarks in order to
reorient themselves when tested in a small room (4 9 6-ft)
but, when tested in a larger room, double in size (8 9 12-
ft), the same children performed comparably to adults.
Hence, the spatial scale of the environment could be crucial
to the children’s ability to deal with the available sources
of information.
Human adults also displayed a similar pattern of
behaviour (Ratliff and Newcombe 2007): trained in either a
small or a large rectangular room with a feature landmark
along one of the walls and then tested after a displacement
of the visual cue only (i.e., while maintaining the room
constant in size), subjects rely on the geometric layout in
the small but not in the large room. In the larger environ-
ment they preferred to use the landmark information.
Moreover, when displaced from a small to a large room or
vice versa and again after the shift of the landmark, sub-
jects preferred to rely on the landmark even in the small
environment demonstrating evidence of an effect of expe-
rience in dealing with featural information.
Comparative studies on the effects of the size of the
enclosure on reorientation in non-human species revealed a
complex pattern of results (see Table 1 for a summary).
Sovrano et al. (2005) tested fish, redtail splitfins (Xenotoca
eiseni), both in a large (31 cm long 9 14 cm wide 9 16
cm high) and in a small (15 9 7 9 16 cm) tank with dis-
tinctively coloured walls that provided featural
information. Fish proved identically able to combine geo-
metric and featural information in both conditions;
however, when displaced from a large to a small tank, or
vice versa, from training to test, they tended to make rel-
atively more errors based on geometric information when
transfer occurred from a small to a large space, and to make
relatively more errors based on landmark information when
transfer occurred from a large to a small space.
Similarly to fish, domestic chicks (Gallus gallus) trained
in a large (70 cm long 9 35 cm wide 9 40 cm high) or in
a small (17.5 9 35 9 40 cm) enclosure showed no dif-
ferences in their reorientation ability, being capable in
successfully conjoining geometry and non-geometry in
either the large or the small arena. However, differently
from fish, chicks did not show any difference in the dis-
tribution of geometric and non-geometric errors when
displaced form the large to the small enclosure or vice
versa (Vallortigara et al. 2005). Only when the chicks were
tested after an affine transformation (i.e., an alteration of
the relations between different information that puts in
conflict geometric and non-geometric information) they
made more choices based on geometry when in the small
enclosure than when in the large enclosure, suggesting that
the use (or the primacy in the use) of geometric and non-
geometric information varies also in this species depending
on the size of the experimental space.
The thread running through all these species, from
humans to fish, seems to be that there is a general ten-
dency to rely more on geometry in smaller than in larger
environments. However, a qualification is needed: the
above mentioned results obtained with chicks refer to a
version of the paradigm in which non-geometrical cues
were supplied as discrete panels placed in correspondence
to every corner (as in the original test with rats), whereas
the children and fish were tested in the blue wall version
of the task (i.e., with one of the walls of the enclosure
made distinctive by use of a different colour). It could be
that in these circumstances chicks were representing space
on the basis of the arrangement of the discrete elements
themselves (the panels at the corners) and, if so, that the
different behaviour displayed in the large and in the small
enclosure was due to an encoding of the landmarks
(panels) at the target and of those close to it that happened
to be localized at different distances in large and small
enclosures. In other terms, it could be that the chicks
tended to use the distant panels to relocate the target
position and that this was more or less easy to do in a
small or in a large enclosure. In fact, if the chicks were
considering the target cue together with its nearest
neighbour along the short wall in the small space, after a
displacement to a larger arena they might consider the
target cue only, since the distances are doubled. On the
contrary, after being moved to a smaller arena, a new
panel appears close to the one located at the target posi-
tion. Hence, the affine transformation would change the
target feature in the small space but not in the large one.
Some recent work specifically addressed this issue.
Chiandetti et al. (2007) trained chicks to find food in a
corner of either a small or a large rectangular enclosure. A
distinctive panel was located at each of the four corners of
the enclosures. No differences in the encoding of the
overall arrangement of landmarks were apparent when
chicks were tested for generalization in an enclosure dif-
fering from that of training in size together with a
transformation (affine transformation) that altered the
geometric relations between the target and the shape of the
environment. The results were therefore not in accordance
with the hypothesis that the chicks encoded the target cue
and its nearest neighbour along the short wall in the small
enclosure but not in the large enclosure.
Further experiments tried to disentangle the relative
contribution of geometry and landmark cues in large and
small spaces (Chiandetti et al. 2007). Again, chicks were
trained to find food in a corner of either a small or a large
rectangular enclosure with distinctive panels located at
each of the four corners of the enclosures. After removal of
the panels, chicks tested in the small enclosure showed
better retention of geometrical information than chicks
tested in the large enclosure. In contrast, after changing the
232 Cogn Process (2008) 9:229–238
123
enclosure from a rectangular-shaped to a square-shaped
one while keeping the corner panels the same, chicks tested
in the large enclosure showed better retention of landmark
(panels) information than chicks tested in the small
enclosure. These findings strongly suggest that the primacy
of geometric or landmark information in reorientation tasks
depends on the size of the experimental space also in non-
human species.
Factors influencing a differential use of geometric and
non-geometric information in small and large spaces
A factor that could affect the probability of using non-
geometrical information in large spaces is the proportion of
the subject’s body size with respect to the experimental
space used (see Nadel and Hupbach 2006). In order to better
understand this point, the same animal should be tested at
Table 1 Summary of results coming from experiments addressing enclosures’ size effects on different species
Study Species Feature cue Space Test change Effect
Hermer and Spelke
(1994)
Human adults Blue wall Small – Geometry + feature
18–24 months old
infants
Blue wall Small – Geometry
2 Landmarks Small – Geometry
Learmonth et al.
(2001)
17–24 months old
infants
2 Landmarks Large – Feature
Landmark Large – Geometry + feature
Blue wall Large – Geometry + feature
Learmonth et al.
(2002)
3–4 years old children Blue wall Large or small – Geometry + feature in
large
Geometry in small
5–6 years old children Blue wall Large or small – Geometry + feature in
large
Geometry + feature in
small (younger children use
still geometry only in
small)
Ratliff and
Newcombe (2007)
Human adults Landmark Large or small Landmark shift Feature in large
Geometry in small
Landmark Large or small Size + landmark
shift
Feature in large and in
small (feature in small
more when trained in
large)
Vallortigara et al.
(2005)
Domestic chicks (Gallus
gallus)
Blue wall Large or small Size (from small to
large and
viceversa)
Good generalisation
Panels Large or small Affine
transformation
Feature in large
Feature + metric + sense
in small
Sovrano and
Vallortigara
(2006)
Domestic chicks (Gallus
gallus)
Blue wall Large or small Affine
transformation
Feature + sense in large
Metric + sense in small
Chiandetti et al.
(2007)
Domestic chicks (
Gallus
gallus)
Panels Large or small Size + affine
transformation
Feature
Removal of panels Geometry (more in small)
Removal of metric Feature (more in large)
Sovrano et al. (2005) Redtail splitfins
(Xenotoca eiseni)
Blue wall Large or small Size (from small to
large and vice
versa)
Good generalisation (more
geometric errors from
small to large; featural
errors from large to
small)
Sovrano et al. (2007) Redtail splitfins
(Xenotoca eiseni)
Blue wall Large or small Affine
transformation
Feature + sense in large
Metric + sense in small
(but also just metric)
Cogn Process (2008) 9:229–238 233
123
different developmental stages while maintaining the same
environmental size; of course, this is not possible since
developmental factors themselves could be sufficient to
explain differences (if any) acting like confounding vari-
ables. However, if we focus on the white-walls task,
different test enclosures were used either across species or
in the same species without affecting the performance. Rats,
for instance, were tested in a chamber that was 120 cm
long 9 60 cm wide (Cheng 1986); domestic chicks (not
much bigger than rats at the age tested) performed the task
in an enclosure of similar size (Vallortigara et al. 1990)as
well as in a smaller one, sized 70 cm long 9 35 cm wide
(Vallortigara et al. 2004; Chiandetti et al. 2007). Hence, no
differences were observed in manipulating geometry.
A second notable element is the size of the featural cue
itself. It may play a crucial role in reorientation perfor-
mance, especially when this non-geometrical information
is provided by a coloured wall, the absolute size of which
changes accordingly in a small or in a large room (Shus-
terman and Spelke 2005). It cannot be ruled out that, when
perceived as being too close by the subject, a coloured wall
may be not properly considered and therefore not pro-
cessed as a cue. Nadel and Hupbach (2006) claimed that
animals prefer to rely upon distal rather than proximal cues
as landmarks for navigation (see also Gallistel 1990). From
this point of view, in small spaces where featural cues end
up located close to the subject, non-geometric information
could be treated as proximal information; in larger spaces,
in contrast, featural cues could be perceived as far-located,
thus increasing the probability for them to be used as distal
information.
This makes sense ecologically since, when navigating in
natural environments, it is more reliable to rely on distal
salient characteristics that do not disappear quickly rather
than on local proximal cues, which are subject to changes
in their relative position while the animal moves in the
environment (Parron et al. 2004). Hence, there may be a
sort of predisposition to treat non-geometric features as
orienting cues (landmarks) only when they are perceived as
far away, but not when they are in close proximity (Nadel
and Hupbach 2006). It is an interesting question in this
regard whether ‘‘absolute size’’ of the wall is important, or
rather if proximity to the subject affects the evaluation,
since in the usual setup a larger room has a wall that is both
greater in absolute size and further away from the subject,
these factors are not the same (and in fact could be
experimentally dissociated).
Recently, Learmonth et al. (2008) tested children of
different ages (from 3 to 6 years old) in various conditions
with the aim of clarifying the role of distal and proximal
landmark use in environments of different sizes. The room
was a rectangular 8 9 11-ft space with one coloured wall;
an inner enclosure sized 4 9 6-ft and 18 inches high was
located in the centre of the room. When the target was
located in the inner enclosure, only 6-year old children
found the correct location whereas younger children relied
only on geometry (confirming previous findings on the
ability of older children to manage correctly all the avail-
able information). When the target was hidden in the outer
room, with the children still confined in the inner rectan-
gular space, 4- as well as 5-year-old infants proved able to
combine geometry and non-geometry to find the goal
object whlist 3-year-olds again chose either of the two
congruent locations (Learmonth et al., in press). This pat-
tern of results is likely to be explained on the basis of a
developmental progression in the use of proximal cues as
informative ones.
Data on the importance of distal cues also come from
neurobiological experiments (Zugaro et al. 2004). Rats
underwent tests in an enclosure in which orienting cues
were provided by the presence of a foreground (proximal)
and a background (distal) card. Electrophysiological
recordings were collected under either continuous light or
stroboscopic light, a condition in which, by disturbing
some dynamic visual processes (such as motion parallax
and optic flow), rats were unable to refer to distal cards.
Responses of head-direction cells appeared to be dependent
on the information conveyed by distal cues, which are
more stable when the animal moves about.
The hypothesis that organisms are prepared to use only
distant featural information as landmarks for reorientation
(Wang and Spelke 2002; Spelke 2003; Hupbach and Nadel
2005) meets, however, with difficulties. One problem with
this view is that, given the evidence of a primacy of geo-
metric information over non-geometric information (see
Cheng and Newcombe 2005 for a review), the basic issue is
not to explain why organisms do not use featural infor-
mation in small spaces (they could do that simply because
of the primacy of geometric information), but rather to
explain why they do not continue to use geometric infor-
mation even when tested in large spaces. This is
particularly intriguing, because it has been usually main-
tained that large-scale geometric information may provide
more stable and reliable cues than local environmental
features such as landmarks (Cheng and Newcombe 2005).
Moreover, species-specific differences in dealing with
the overall sources of information should be considered.
Conceivably, reorienting making use simultaneously of
metric (long–short), features (white–blue) and sense (left–
right) information could be costly: it may become difficult
to handle all this information together, at least in non-
human species, and therefore some of them could be sep-
arately considered with a preserved possibility to reorient
successfully.
Sovrano and Vallortigara (2006) produced evidence
suggesting that sense for left–right distinction could be
234 Cogn Process (2008) 9:229–238
123
separately associated with geometry and landmark infor-
mation depending on the spatial scale of the environment.
Consider the blue wall version of the task, and the condi-
tion in which the correct position (faced by the chick, in
Fig. 2) is properly defined by having, while facing it, a blue
short wall on the right and a long white wall on the left.
The encoding process that could be used here may rely
either only on sense (left–right) and metric (long–short)
thus having a definition like ‘‘a short wall on the right and a
long wall on the left’’, or only on sense and the non-geo-
metric cue thus having a definition like ‘‘a blue wall on the
right and a white wall on the left’’.
When chicks were trained in either a large or a small
enclosure and then tested in enclosures of the same size as
used during training, after a blue wall dislocation (e.g.,
with the blue wall on the short wall during training and on
the long wall at test) they showed a different linkage of
sense information with either metric or landmark infor-
mation depending on the spatial scale of the environment.
That is, in small spaces chicks linked sense with metric
properties of surfaces, in large spaces they linked sense
with local landmark cues (Sovrano and Vallortigara 2006;
and see also Sovrano et al. 2007 for similar results with
fish).
This is shown in Fig. 2. Assuming that visual analysis
of a corner (for instance by head-direction cells) occurs at
a fixed distance from the animal, then in a small envi-
ronment (Fig. 2, right) the available information provided
by complete scanning of the length of the surfaces may
provide a reliable source of information for spatial reori-
entation. Thus, animals may rely on an association
between the metric properties of the surfaces and the sense
(the correct corner has a short wall on the right and a long
wall on the left). In a large environment, however, scan-
ning of the full spatial extent of the surfaces is prevented
(Fig. 2, left). Thus, the animals must rely on an associa-
tion between the featural properties of the surfaces and the
sense (the correct corner has a blue feature on the right
and a white feature on the left). According to Sovrano and
Vallortigara (2006) such different associations depend on
what the animal is pondering in environments of different
spatial scale.
This hypothesis is not necessarily in contrast with claims
for a preference of using distal rather than proximal cues
for reorientation, but provides a more precise account of
the overall pattern of findings with the rectangular task in
large and small enclosures in a variety of species.
Further support for this view comes from results
obtained with people with Williams syndrome, a genetic
deficit that severely compromises spatial representations
while relatively preserving language (e.g. Brown et al.
2003): these patients are good performers when tested in
the blue wall task thus proving able to conjoin a metric
property with the left–right sense whereas they fail in the
white-wall version, i.e., fail in using geometry alone
(Landau et al. 2006). Obviously this would imply that in
the blue wall task these subjects would perform differently
when the target was located between two same-colour
corners (in which only metric information can be used) or
between two differently coloured corners (in which colour
plus sense can be used as an alternative to use of metric
plus sense). No evidence for such a difference is currently
available. Interestingly, however, Nardini et al. (2008)
reported recently that children of 18–24 months old can
reorient using the left–right sense of coloured landmarks
(and see also Burgess 2006).
Concluding remarks
Many different species have been shown to be able to
reorient themselves encoding and remembering the geo-
metrical information provided by the metric arrangements
of the walls of a rectangular enclosure together with
available non-geometric cues. However, differences in the
relative reliance on geometric and non-geometric cues have
been observed when animals were tested in enclosures of
different sizes.
The results discussed in this review bring forward evi-
dence that animals prefer to use geometric information
when in a small environment and prefer to rely on land-
mark cues when in a large environment.
Fig. 2 Photograph layout of encoding occurring in a large (left) and a
small (right) rectangular enclosures with a blue wall
Cogn Process (2008) 9:229–238 235
123
Given that there is substantial agreement in arguing for a
prominent role of geometry in natural environments (see
Introduction) due to its stable and invariant characteristics
across time, it is not difficult to believe that vertebrates are
somehow innately predisposed to manage geometry for
successful navigation (Chiandetti and Vallortigara 2008;
Vallortigara and Sovrano 2002). Hence, a hierarchical way
to process different kinds of information resembles the
hierarchical structure of the environment.
When we think of a natural environment, it is not dif-
ficult to note that usually several redundant cues are
available and that if one organism simply relies on just one
at a time the strategy could prove successful. Such a pattern
of reliance (one at a time) may directly reflect the path of
evolution (Simon 1969) with an addition of new modules
rather than a change in the old ones (Shettleworth 1998).
In order to unravel how they are weighted, different cues
can be placed in conflict so that one cue indicates one goal
location and the other cue another goal location. The rel-
ative weightings of different sources of information may
change with different conditions and it can happen that the
animal resorts to use one whilst disregarding another. Thus,
some cues may obtain a primacy in processing and hence
they may provide a context for subsequent analyses (see
also Shettleworth 1998).
However, the unresolved question is, why do animals
resort to using geometry in a small environment but prefer
landmarks in larger ones. A challenging hypothesis comes
from the possibility that there is a different association
between metric, sense and landmarks cues as suggested by
Sovrano and Vallortigara (2006). When located in a small
enclosure an organism has the metric layout of the surfaces
around it available to it, thus adding the sense for left–right
discrimination it has a reliable source of information for
spatial reorientation. When an organism is in a larger
enclosure, estimation of the lengths of extended surfaces
would require costly scanning or direct movements back
and forward: this could lead it to prefer to associate a
featural property immediately available in the vicinity with
the left–right sense, thus discarding use of geometry.
Such different associations could be expected since they
convey more reliable information to an organism on the
basis of visual (or other sensory) scanning of environments
of different spatial scale.
However, what does reliable really mean here with
respect to size? If we assume that reliability in large as well
as small environments is directly linked to the source
of information that conveys the most information for the
least effort (i.e., efficient information) and it is also the
most stable, the type of information on which there is a
reliance can change according to the size of the environ-
ment. In a small environment things change rapidly as the
animal moves; in this case, geometry is the most stable
information whereas local information is difficult to com-
pute. In a large environment, the features are less likely to
change: while moving, they require a long time to disap-
pear from the visual field. In this sense, landmark cues are
stable and informative. Now, it is useful to distinguish
between directional and positional information. All distal
information allows great efficiency in guiding navigation
because—as discussed in the previous sections—it pro-
vides stable directional and useful positional information.
Proximal cues may have more precise positional informa-
tion but offer a much reduced directional information.
Moreover, motion parallax effects and large changes in
visual signals are high for proximal landmarks, therefore it
would probably be simpler relying on distal landmarks in a
large environment and it would probably be less difficult
using metric differences in a small environment, in the
sense that each kind of information acquires different
values of predictive strength as a function of their
usefulness.
Thus, an alternative explanation can be put forward on
the basis that two different mechanisms are involved, one
for distal and one for proximal cues’ use, as suggested from
neurobiological works, at least with respect to rats (Parron
et al. 2004). Specifically, it has been shown that entorhinal
cortex lesioned rats were compromised in using distal
landmarks but not proximal ones (Parron et al. 2004). Of
course, the role of the entorhinal cortex in navigation is far
from being clear, and it is not the aim of this review to
discuss it. However, the authors suggest that enthorinal-
hippocampal circuitry processes the distal cues whereas the
parietal-hippocampal circuitry accounts for proximal cues.
Moreover, single unit recording studies on place cells
(Cressant et al. 1997) as well as on direction cells (Zugaro
et al. 2001) seem to confirm a dissociation between distal-
proximal landmark processing.
What exactly happens while an animal is reorienting in
environments of different spatial scale is of course still
matter of debate (see Lee et al. 2006). The two alternative
views suggested in this review are not in contrast. From the
behavioural side, there should be another (unverified)
condition replicating Chiandetti et al. (2007) condition
(with both the change in size and the affine transformation
from training to test) but with the blue wall; in this con-
dition it should be expected a stronger effect on different
linkage among separate kind of information on the basis of
the experimental space size used. From the neural side,
data coming from neurobiological studies will maybe bring
some clarifications on the way different sources of infor-
mation are encoded and processed before the overt choice
of the organism.
Acknowledgments This work was supported by grants MIUR Cofin
2004, 2004070353_002 ‘‘Intellat’’ and MIPAF ‘‘Benolat’’ (to GV).
236 Cogn Process (2008) 9:229–238
123
We would like to thank two anonymous referees for their suggestions
on a previous version of this manuscript.
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