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97

DETOUR AND SHORTCUT ABILITIES IN SEVERAL SPECIES OF MAMMALS.

Nicole CHAPUIS

Department of Animal Psychology


Institut of Neurophysiology and Psychophysiology
CNRS-INP 9
31, chemin Joseph-Aiguier
13402 Marseille Cedex 9
FRANCE

This study is concerned with t\vO properties of cognitive I'lapping. The


first is plasticity, by which I mean the ability of an animal to
reorganize its previous experience of a given situation. Thus, for
example, when modifications are introduced into a faDiliar spatial task,
some anil'lals can find original solutions ; they do not become lost and
they can still reach the goal. The second property is optimalization. It
entails the choice and the planning of the best adapted solution : for
example, taking the most direct of several possible ways to reach a goal.

In addition, I am particularly concerned with the question of how


cognitive maps are built up, and what strategies of response various
species of mammals employ here. My detour and shortcut experiments were
conducted in "free" situations, Le. in openfields in which no route was
laid out, so animals could choose-b;tween various available ways to reach
the goal. For the DOSt part, they were conducted outdoors, in a natural
landscape. The subjects were dogs, cats, hamsters and horses.

Their response strategies were examined in relation to previous


information collected by animals in their own displacements that is to
say kinesthetic and visual (etc.) information originated from the
landscape.

DETOUR EXPERIMENTS

The first set of experiments consisted of detour tasks, in which


animals had to circumvent an obstacle, or to move away from the goal in
order to reach it. Such tasks have long been, since Kohler (1925), to
constitute tests of insight learning. Detour abilities have been
del'lonstrated in several species of mamoals and also in human infants by
Piaget (1937) and by Lockman (this book, vol. 2). Several factors have
been shown to affect this ability. Among them are whether or not the goal
is visible from the choice-point, the nature of the cues whereby the goal
can be located, and the length, the number and the complexity of paths
leading to the goal.

Our experiments were done on cats (Poucet, Thinus-Blanc and Chapuis,


1983), dogs (Chapuis, Thinus-Blanc and Poucet, 1983), and on horses with
Tournadre (note 2). In an openfield, one or several screens were placed
between a starting-point and a baited feeding-bowl (figure 1).

P. Ellen et al. (eds.), Cognitive Processes and Spatial Orientation in Animal and Man
© Martinus Nijhoff Publishers, Dordrecht 1987
98

Arrangement of the screen(s) was devised so as to leave two routes ;


these routes differing in their respective lengths (short or long) and/or
angular deviation (narrow or wide) related to the starting-point - goal
axis. Two conditions were used : in the first one, the goal could be seen
from the choice-point (transparent screen), vlhereas in the second, it
could not (opaque screen). The aim was to assess the part played by these
two factors length and angular deviation in a visually guided
condition (goal visible) and in an "inferential" condition (goal hidden).

I
G II
"{---?\ II
rr
\ :\ II
\ i \ G! I
\ : \ _"". I
\ , \
\ """",""" ," ~
"*<?
~,
\: \ /~
~ \
~*~ 5
s

I :IT ill

Figure 1. Overviews of the apparatus, with location of the starting-point


(S), feeding-bowl (G), screen (solid) and rOUteS (dashes) in the four
situations. Adapted from Chapuis et al., 1983 copyright The
Experimental Psychology Society. -- --

Four experimental situations were presented to the animals, as shown


on figure 1, in the two conditions: goal visible and goal hidden.

SITUATION I
Angular deviations and lengths were both relevant and convergent,
i.e. the shorter way was the less divergent and the longer way was the
more divergent from the goal direction.

SITUATION II
The two ways differed by the angular deviations but were siElilar in
length.

SITUATION III
The angular deviations were equal but the lengths were different.

SITUATION IV
Angular deviations and lengths were both relevant but opposed : the
shortest way was the more di vergent and the longer way \13s the less
divergent.
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Subjects could reach the feeding-bowl by either of the t\VO ways and
were allo\.ed to eat food regardless of the way they had gone. Prior to
the beginning of testing, the animals were trained to run froI!1 a
starting-point straight to a feeding-bO\·,l which was not blocked by a
screen. Following this initial training period, the screen was
introduced. Subjects were tested for eight days with twelve trials per
day. After six daily trials, the location of the screen and the feeding
bowl was changed symmetrically about the axis defined by the
starting-point and the goal. Each way (as defined by the parameters in
figure 1) appeared equally often to the right and the left of the
symmetrical axis. At the beginning of the daily test, before the first
trial, and after the change of the location of the screens (before the
seventh trial), the animals were led from the starting point to the goal
along each of the two routes in order to show them the place of the
feeding-bowl behind the screen.

PARAMETERS Angular deviation Angular deviation Length Angular deviation


SPECIES STUDIES and length SITUATION II SITUATION I II versus angle
SITUATION I SITUATION IV
STUDIED
WAY less divergent 1ess di vergent shorter way shorter and more
CHOSEN and shorter way way divergent way
---
C visible goal 81.25 % 70 % 65 % 51.25 %
A condition lU U II no preference
T
S 93.75 %Ul
hidden goal 75 % (w/out distal cues: 83.75 % 85 %
condition 62.5 %) l
-_ ... -- ---- -- ----
Ul III III
.------------- -- ---_ ... ----- -- -- ---- - ......
·-------~ ___________ a _________________

H visible 81.10 % 72.2 % 76 % 58 %


o l+ no preference
R
S
E hidden 92 % 66 % 64 % 63 %
-S------ ------------ -----!!! _________ .______ !L ___________ . ______! __________ !lQ J!r~f~!:~!l!:!L_
D visible 95 % 71. 7 % 53.3 % 50 %
o III + no preference no preference
G
S hidden 96.7 % 45 % 75 % 68.3 %
III no preference + +

(Student's t test + P < .10 l P < .05 l+ P < .02 II P < .01 III P < .001)

TABLE I. Percentage of trials on which the animals chose various


different ways of circumventing an obstacle.
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The general procedure of the experiment IVas the same for the three
species. But, of course, the test-situation itself IVas modified (for
example in size) to fit each species. The experimental frame also IVas
different since the eight cats IVere tested in a large indoor room IVhile
the six dogs and the nine horses IVere tested outdoors : dogs in a large
meadolV and horses in a paddock.

As Table I sholVs, in situation I cats selected the shorter and less


divergent path, IVhether or not the goal could be seen. In situation II,
where the angular deviation from the goal IVas the only factor, cats used
the available parameter in the both conditions. TIVo tests were done with
the opaque screen. In the first one, the visual cues of the room were
available while in the second one they were not available because of
black curtains on the IValls. The trend to choose the less divergent IVay
was greatly reduced by reference to the scores obtained previously. So
the visual cues seemed to be used by cats to localize the place of the
goal hidden behind the opaque screen. In situation III, the shorter path
was preferred in the both conditions although a significant difference
appeared between the results in the two conditions. Lastly, in the
conflictual situation situation IV a preference appeared for the
shorter way only when the goal was hidden. So, both angular deviation and
length were relevant parameters for this experiment when they IVere alone
or associated. \~hen they were opposed, the shorter way could not be
selected when the goal IVas perceived from the choice point. Thus the
visible goal appeared to act as a "perceptual anchor" and to eliminate
the use of distance information.

The results of horses were similar to cats' responses except in


situation IV where they did not show any preference even when the goal
was hidden.

In contrast, the behavior of dogs was a bit different since, on the


whole, opposed behavioral strategies IVere used by the same subjects : the
dogs preferred the shorter way when the goal was hidden but not when it
was visible. They more often chose the less divergent way when the goal
was visible than when it IVas not. Their performance in situation II, in
the hidden goal condition, seems surprising since shortcut results (to be
presented later) have sholVn that dogs were able to evaluate accurately
the direction of a non-visible goal. The difference between dogs and cats
(and horses) can most likely be attributed to differences in the number
of environmental stimuli available to the animals. The cats IVere tested
in an empty room with differentiated walls ; a large window was at South,
a blind wall at West, a small window at North, and a door at East. The
horses were tested in an area I,hich has a fence and several buildings
around it as stimuli. The field in which the dogs were tested was devoid
of prominent visual distal landmarks. To assess accurately the correct
position of the hidden goal was rather complex for dogs. It is likely to
imply the use of an egocentric reference framework, the body axis, which
was oriented towards the middle of the screen, at the starting-point, to
know the position of the goal (left of right) with regard to this body
axis. The lack of preference of dogs in situation III when the goal could
be seen in the middle behind the screen support our previously expressed
idea of a "perceptual anchor".
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Thus, two classes of mechanisms appeared to be involved when the goal


was hidden and when it was visible. When the location of the hidden goal
must be remembered, animals were able to take ircto account and integrate
the information collected during the prior exploratory runs in order to
plan the shortest route. The direct visual perception of the goal, on the
other hand, allowed the animals to use a guidance strategy that led the
subject to reduce the distance bet~een itself and the goal as quickly as
possible, which, in this case, was the simplest str2tesy.

SHORTCUT EXPERIMENTS

The second set of studies investigated the animals' ability to take


shortcuts. The first experiment (Chapuis and Varlet, note 1), was carried
out with seven dogs in a 3 hectare meadow. The animals' task was to find
meat hidden at two points (A and B), when released from a third (D)
(figure 2). On each trial, the dog had been taken previously on a leash
to the two baited points by a different path that led indirectly from A
to B via D, i.e. along the DA-AD-DB-BD route. They were shown the food at
points A and Bwithout being allowed to eat it.

Ba

B6·········· .. · .... A6 ~B4

B5

Figure 2. One standard experiment situation. Course of one trial. D


corresponds to the starting-point, A and B to food-points. DA-AD-DB-BD
vectors (solid) represent exploratory runs (dogs on the leash) and AB
segment (dotted) the optimally oriented task run ("direct shortcut").
From Chapuis and Varlet, note 1 ; copyright The Experimental Psychology
Society.
Figure 3. Experimental set-up showing the arrangement of the eight
standard situations (DA B) on one field (ground HET). From Chapuis and
Varlet, note 1 ; copyrigh~ The Experimental Psychology Society.
lO2

The experiment was carried out in two different fields, one nearly
uniform - it was covered mainly with thyme - and the other heterogeneous,
containing visual features such as bushes, puddles and trails. Within
each field, each pair of baited points was changed eight times to
different A and B places as shown in figure 3. In 96 % of the 224 trials,
the dogs took a shortcut between the baited points rather than returning
to the start.

A qualitative analysis of the characteristics of the shortcut


trajectory i:1duced me to distinguish between the "direct shortcut"
(strategy I) which consisted of a straight line between A and B and the
second and the third classes, which included trajectories making a
smaller or a wider angle than the angles DAB or DBA ; so the path was
either inside the triangle DAB, and corresponded to a strategy II or
"inside shortcut", or the path was outside DAB, and corresponded to a
strategy III or "outside shortcut". Strategy IV consisted of following
the indirect route ADB (or BDA) which had been shown to the dog in the
exploration phase. As can be seen in figure 4, the dogs used strategy I
more frequently in the uniform than in the heterogeneous field.

STRATEGY

Figure 4. Frequency of different classes of responses (: S.E.M.) produced


by the seven subjects tested 16 times on each type of field,
heterogeneous (HET) and uniform (UNI). I : direct shortcut, II : inside
shortcut, III outside shortcut, IV : return. From Chapuis and Varlet,
note 1 ; copyright The Experimental Psychology Society.

In the litter field, the animals had a greater tendency to use an


"inside shortcut" (strategy II), heading for a point which intersected
the path from the start to the second baited point rather than going
directly to the baited point or returning to the start.
103

The shortcut responses presented in figure 5 are an example of the


trajectories of one subject on the two grounds, heterogeneous and
uniform. With this dog, shortcut responses were distributed only between
strategies I and II.

81

88

83 81
11

814

85 813

GROUND HET GROUND UNI

Figure 5. The map of the sixteen trials performed by one subject (number
3) which used the strategy of the "direct shortcut" in situations AlBl'
A1 B3 , B4A4 and ASBS on ground HET and in the eight situations on ground
UNI. Thls subject used the strategy of the "inside shortcut" on the four
other situation~ on ground HET, .:!:..~. AZB Z' BsAs, A6B6 and A7B7. The
numbers noted ln each DA B triangle correspond to the temporal order in
which tests were present~dnto the subject. From Chapuis and Varlet, note
1 ; copyright The Experimental Psychology Society.

The trials in which the shortest path was taken suggest that dogs
are able to evaluate, along a continuum, the direction of an invisible
goal by integrating motor and/or sensory cues ohtained during an early
but indirect visit to the same goal.

On the other hand, I do not think that the use of the inside shortcut
is due only to a misestimation of the direction of the second goal, since
(if such were true) the distribution of exterior and interior shortcuts
would be equivalent. The dogs' behavior could be interpreted as a
compromise between returning to the start and heading for the second
goal. From an adaptive point of view, it could correspond to the pursuit
of safety. Depending on the meaning of the word safety, we can suggest
two clearly related hypotheses. According to the first hypothesis, the
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probability of finding the line DB is greater than the probability of


finding point B if the dog is not certain of the direction of the goal B
when it is at the goal A. According to the second view, in using this
strategy, the dog is able to find known territory by the shortest
possible new way, without moving away from the goal as would be the case
if it had used the path ADB. Observations of human orientation incline
one towards the latter two hypotheses. Studies by Pailhous (1970) have
shown, for example, that when Paris taxi-drivers found themselves in an
unknown area, they did not attempt to drive directly towards their goal,
but took the shortest possible route to reach a well-known major route,
which they used to help them towards their goal.

It was also observed that dogs' shortcut responses were more accurate
in the uniform field than in the heterogeneous one since direct shortcuts
were more often taken than inside shortcuts in the first case and
vice-versa in the second case. To encode the location of the two goals A
and B, the dog must attend either to distal cues, to information obtained
from locomotion (kinesthetic and visuo-kinesthetic cues), or to an
inter-relation between them. In the heterogeneous field, they could of
course also attend to proximal cues during their previous run on the path
DA-AD-DB-BD. Note that no cue was directly related to the goals. So, in
the uniform field, while at goal A, dogs had to decide the most direct
way to take on the basis of the kinesthetic information and/or distal
cues acquired during the previous exploratory run. Conversely, when the
field contained landmarks, they did not need to work out the precise
direction of the second goal B, since they could also use immediate local
landmarks on the path DB.

A complementary experiment was carried out with six naive dogs. Here,
we varied the size of angles DAB and DBA during the experiment. This
experiment was conducted in a heterogeneous field. The results are
consistent with the previous results since dogs took the inside shortcut
in 38 % of the responses, and the direct shortcut only in 21 % of the
cases.

On the other hand, in shortcut experiments with hamsters conducted in


a wheel-maze (Chapuis and Lavergne, 1980), a strategy very similar to the
inside shortcut was observed. We called it "intercepting strategy". It
occurred in the case where the task was too difficult to be solved by a
direct shortcut, that is when the previous exploratory displacement was
very complex and also when no environmental cues were available.

Behavioral constants can thus be found among various species of


mammals ev~n including humans. Even if a species has the ability to
integrate information in order to take the most direct or the shortest
route to a goal, the animals do not necessarily use that ability. They
may prefer to choose the surest route (one with which they never loose
sight of the goal, for instance) or to head for a route to the goal with
which they are already familiar.

All these results show the plasticity of the behavior of the mammals
studied and also point to an optimalization of the responses, since the
subjects succeeded in detour and shortcut tests. The concept of
optimalization needs however to be enlarged beyond its classical
acceptation in orientation studies, which is the taking of the shortest
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way to reach the goal, implying the saving of time and kinesthetic
energy. Factors contributing towards minimization of the risks, such as
the risk of becoming lost or vulnerable to predators, should also be
taken into account. The notion of "least risk" does not always correspond
to the "least effort" principle discussed by Tolman (1932), since the
trajectories of animals in a given task sometimes cannot be analysed only
with physical criteria such as the shortest time or the shortest distance.
In short, \~e can say that in the present study the animals adapted
their behavior depending on their perceptual and cognitive abilities,
their motivations, the variety and complexity of the landscape and
whether or not they were given the possibility of acquiring kinesthetic
information as well as information provided by landmarks.

REFERENCES

Chapuis, N. and Lavergne, F. 1980. Analysis of space by animals.


Visual and proprioceptive cues in complex spatial tasks by hamsters.
Neuroscience Letters Supplement, 5 : 171.

Chapuis, N., Thinus-Blanc, C. and Poucet, B. 1983. Dissociation of


mechanisms involved in dogs' oriented displacements. Quarterly
Journal of experimental Psychology, 35B : 213-219.

Kohler, W. 1925. The mentality of apes. New York Harcourt Brace Co.

Pailhous, J. 1970. La representation de l'espace urbain. L'exemple du


chauffeur de taxi. Paris : Presses Universitaires de France.

Piaget, J. 1937. La construction du reel chez l'enfant. Neuch~tel


Delachaux et Niestle.

Poucet, B., Thinus-Blanc, C. and Chapuis, N. 1983. Route-planning


in cats related to the visibility of the goal. Animal Behaviour,
31 : 594-599.

Tolman, E.C. 1932. Purposive behavior in animals and men. New York
Appleton-Century-Crofts.

Notes
1. Chapuis, N. and Varlet, C. Shorcuts by dogs in natural surroundings.
Quarterly Journal of experimental Psychology (In press).
2. Tournadre, V. 1984. Analyse des parametres impliques dans les
deplacements orientes chez Ie Cheval a travers des experiences
de detour. Memoire de Maitrise de Psychologie, Universite de
Dijon.
106

Acknowledgments

These experiments were carried out on the grounds of the Centre Regional
d'Elevage et de Production d'Animaux de Laboratoire in Rousset, France.
I would like to thank Professor Emil Menzel from the State University of
Nel"l York, and Dr. Charlene Wages, from Georgia State University in
Atlanta for their suggestions on this manuscript.

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