Behavioural Processes 79 (2008) 1927

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Behavioural Processes
journal homepage: www.elsevier.com/locate/behavproc

Chimpanzees fail to plan in an exchange task but

succeed in a tool-using procedure
V. Dufour a,b, , E.H.M. Sterck b,c
a
Ethology Research, Animal Science Department, Biomedical Primate Research Centre, Lange Kleiweg 139, 2288 GJ Rijswijk, The Netherlands
b
Behavioural Biology, Utrecht University, Kruijt Gebouw Room O 211, Padualaan 8, 3584 CH Utrecht, The Netherlands
c
School of Psychology, University of St. Andrews, St. Marys College, South Street, St. Andrews, Fife, Scotland KY16 9JP, United Kingdom

a r t i c l e i n f o a b s t r a c t

Article history: Planning has long been considered a uniquely human capacity. Lately, however, it has been shown that apes
Received 28 November 2007 and a corvid species act now to derive a material future benet. Since primates are highly social animals
Received in revised form 29 March 2008 and their sociality is considered a strong selective force that resulted in complex cognitive capacities,
Accepted 11 April 2008
planning is also expected in social situations. Unfortunately, prompting from social partners cannot be
excluded in a social setting. Therefore, we controlled for this factor by testing the capacity to plan in
Keywords:
chimpanzees using an exchange paradigm, that involves both a material and a social component, and a
Planning
tool-use paradigm, similar to the one used on two other ape species. All chimpanzees failed to plan in
Tool-use
Exchange task
the exchange task, but three individuals showed planning behavior in the tool-use task. Our methods
Cooperation controlled for the fact that chimpanzees were not prompted by the visibility of the reward at the moment
Chimpanzees of planning and also could not repeat a previously acquired routine. The best interpretation for our results
is that chimpanzees can plan. However, planning was limited to the situation where the action to attain
the future benet only depended on a chimpanzees own behavior.
2008 Elsevier B.V. All rights reserved.

Whether humans are the sole species that evolved episodic
memory, the capacity to mentally travel in time, is currently sub-
ject of a heated debate (Schwartz and Evans, 2001; Clayton et al.,
2003a,b; Roberts, 2002; Suddendorf and Busby, 2003a,b). Many
researchers (Roberts, 2002; Suddendorf and Busby, 2003a) claim
that animals have no recollections of personal events and con-
sequently are not able to plan for future needs. Instead, it has
been suggested that animals are stuck in time (Roberts, 2002).
Animals generally favor the immediate reward (Richards et al.,
1997; Tobin et al., 1996), when exposed to one small immediate
and one larger but delayed reward. Silverberg et al. (1998) noted
that chimpanzees (Pan troglodytes) were unable to stock fruits in
advance, thus exhibiting temporal myopia the inability to antic-
ipate a future need . Yet in a non-primate species, Clayton and
Dickinson (1998) showed that scrub jays (Aphelocoma californica),
a food hoarding bird species, possess episodic-like memory of a
past event, since they integrate information on what-where-when
to recover cached food (Clayton et al., 2001). Moreover, scrub jays
react to the potential of future pilfering by reaching the food (Dally
et al., 2006) and future planning was suggested as an explanation
for food caching behavior in the evening before a predictable lack
Corresponding author. Tel.: +44 7954 989813; fax: +44 1334 463042.
E-mail address: vmd1@st-andrews.ac.uk (V. Dufour).
of breakfast the next morning (Raby et al., 2007). Also primates
show future-oriented behavior. Monkeys going to distant food sites
choose the appropriate route minutes in advance (Cebus apella,
Janson, 1998). Captive chimpanzees spontaneously save tokens,
indicating anticipation of a delayed opportunity to exchange them
(Sousa and Matsuzawa, 2001). Recent work (Dufour et al., 2007;
Beran and Evans, 2006) shows that chimpanzees can postpone an
expected reward for a time scale measured in minutes, not seconds.
In both common chimpanzees and bonobos, individuals trained to
use symbols were able to announce intended action and travel des-
tination plans (Menzel et al., 2002). However, only one study has yet
experimentally shown that apes performed future-oriented behav-
ior that was not driven by current need, a necessary condition for
planning (Mulcahy and Call, 2006). In this study, orangutans (Pongo
pygmaeus) and bonobos (Pan paniscus) collected, saved and trans-
ported tools to use them later. Individuals of both species could plan
for a predictable event 1 h, and even one night, ahead. It has, how-
ever, been argued that prompting, the food was visible at the time
of tool collection, or learning from the rst task to solve the sec-
ond task may also explain these results (Suddendorf and Corballis,
2007).
Advanced cognitive capacities, such as future planning, may
have evolved in primates to cope with a complex environment.
This complexity may concern their social environment (Byrne and
Whiten, 1988), extractive foraging requiring tool use (Reader and

0376-6357/$ see front matter 2008 Elsevier B.V. All rights reserved.
doi:10.1016/j.beproc.2008.04.003
20 V. Dufour, E.H.M. Sterck / Behavioural Processes 79 (2008) 1927
Laland, 2001) or a combination of the two (Reader and Laland,
2002). Anecdotal evidence on chimpanzees suggests that they are
able to plan for future material needs. Wild chimpanzees travel
over hundreds of meters in dense vegetation to pick up and trans-
port tools necessary to crack nuts (Boesch and Boesch, 1984).
Chimpanzees may also plan for future social needs. They seem to
maintain a high dominance rank through tactical social maneu-
vering (de Waal, 1982). Moreover, chimpanzees that groom in the
morning have a higher chance to obtain a share of food from
their grooming partner in the afternoon (de Waal, 1997). However,
interpreting these observations as evidence of planning remains
speculative since other, lower-level, explanations are possible. For
example, chimpanzees tool transport may result from the cur-
rent motivational state of the individual (Suddendorf and Corballis,
1997). Moreover, in a cooperative context chimpanzees fail to show
other regarding behavior (Jensen et al., 2006; Silk et al., 2005) or
to understand cues givens by others who communicate coopera-
tive intentions (Hare and Tomasello, 2004). As suggested by Byne
(2007), cooperation observed in chimpanzees may not result from a
calculation of future benets (planning), but from their inclination
to favor those with whom cooperation has been successful before.
Herrmann and Tomasello (2006) have shown that chimpanzees
express more cognitive skills when in a competitive context rather
than in a cooperative one: chimpanzees deduce from a commu-
nicative gesture from a competitor toward a bucket that the food
must be hidden there. A similar gesture made by a cooperative part-
ner with the intent to communicate the location of the food is not
understood. Cognitive skills in a cooperative context may be con-
strained by the high level of competition in this species (Hare and
Tomasello, 2004; Hare et al., 2007). If this is the case, planning for
the future involving cooperation may be a more challenging task
than planning for a future activity where success only depends on
ones own behavior.
We aimed to test the capacity to plan in chimpanzees. Most tests
of planning in animals concern a future material benet and do not
involve uncertainty related to other factors such as a cooperative
partner. In the present study we tested for chimpanzees planning
capacities involving material benets based on a cooperative inter-
action (Part 1), and planning capacities involving material benets
only (Part 2).
1. Study 1: planning to selectively collect items for later
exchange
To be qualied as planning, a behavior must be performed in
the present without being driven by current need (Tulving, 1983)
or by current cues to future benets (Suddendorf, 2006). Tulving
(2005) proposed the spoon-test to test for planning in animals. In
this test, a girl was disappointed because she could not get desert
at a party since she did not bring her own spoon. The crucial test
for planning capacities is to check whether this memory of disap-
pointment will lead her to bring a spoon to the next party. In this
study, we devised an analogue of the spoon test (Tulving, 2005), to
assess chimpanzees capacity to plan for a future benet involving
a cooperative interaction. Unfortunately, purely social interactions
cannot be used to test future directed behavior, since the antici-
pated future event will always involve other individuals and direct
reactions to the others cannot be ruled out. Therefore, we tested
chimpanzees capacities in an exchange paradigm, where the coop-
erative individual was represented by a human partner. While the
human partner could be considered as a social tool, failure may be
considered from the subjects point of view as a failure in the part-
ner behavior, and not his own. From this point of view it involves a
social component that the subject cannot control. In this paradigm,
a food reward could be obtained from a human partner upon the
return of a specic object at a specic time of the day. To succeed in
this task, chimpanzees rst had to collect earlier in the day a specic
category of object among three available for a limited time inter-
val, and return with it in the test compartment where the exchange
activity would take place with the human partner.
1.1. Methods
1.1.1. Subjects
Two groups of chimpanzees from the Biomedical Primate
Research Centres colony participated to this study. They were
housed in a similar enclosure with the same facilities and daily rou-
tine. Group 1 was composed of two males and three females aged
from 19 to 23 years. Group 2 was composed of two females and four
males aged from 12 to 15 years. Each group was housed in its com-
munal room (a 40 m2 room with elevated sitting devices) to which
individual feeding compartments were permanently connected.
These compartments were used as individual testing compart-
ments. One individual in group 1 never learned how to use objects
and was therefore not included in the study. The study was designed
to provide enrichment without disturbing the daily routine of the
groups. Participants were given the choice to come or not and where
always rewarded with a small treat at the end of the session, what-
ever their level of participation to the task. Both groups had never
been observed for ethological purposes before. They had never been
trained for or involved in cognitive tasks. The chimpanzees were not
reported to have been trained to exchange with humans before.
1.1.2. Training
Prior to the tasks, the chimpanzees were all invited to enter their
individual compartment, a familiar procedure. For the exchange
task, chimpanzees were trained to return an object to the human
partner to obtain a food reward. The object to trade was a color-
ful plastic straw that was easy to insert through the front of the
compartment and was unknown to all subjects. The human part-
ner (V.D.) proceeded as follows: she begged the chimpanzee with
the palm up presented in front of her/him, holding a peanut in the
other hand, and asked verbally for the return of the token (terug
the Dutch equivalent for back) 12 times in a row. After a successful
exchange, the chimpanzee was given 15 s to eat the peanut. Chim-
panzees learned to give the straw back with a mean of 4.7 sessions
of 12 trials each (range 117 sessions). Once all individuals mas-
tered the exchange task, this task was given by the partner as a
daily activity for 20 consecutive daily sessions at 13:10 every after-
noon. This insured predictability of the activity and also insured
that the human partner was associated with the exchange task.
The human partner had at the start of this study been known to
the chimpanzees for half a year and she was only involved in neu-
tral or positive interactions with the animals. The chimpanzees had
known the keepers from at least 5 years.
To determine whether the chimpanzees planned for the
exchange task at hand or were interested in just any object, other
objects were present as distracters at the time of testing. These
objects, like the straws used in the exchange activity, were also
associated with a desired food, a specic function, and a specic
time of the day. The chimpanzees were provided a natural wooden
branch that was to be used in a dipping task. The chimpanzees
were trained to dip it in a box full of honey that was held in front of
them by a keeper. Since the dipping task was part of their weekly
enrichment, only 3 sessions of 3 min per day and per individual
were required to insure that all the individuals performed the task.
The third object consisted in a sharp wooden stick that was used in
a picking task. The chimpanzees were trained to retrieve, by pick-
ing into them, orange pieces from a plastic box held in front of the
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