e
r
Pla
able not only to react to what others are doing but also to
Review
others in many situations based on past experience (and
perhaps specialized cognitive adaptations), but they do not
which chimpanzees begged indiscriminately from humans
facing them and others with buckets over their heads (see
also Ref. [7]). There was also one negative study on chim-
panzees’ understanding of false beliefs [8].
All of these data led Tomasello and Call [1] to the
general conclusion that chimpanzees and other non-
human primates do not understand the psychological
states of others. That is, they can predict the actions of
anticipate what theywill do. Oneway of accomplishing this
is by observing what others do in particular situations and
deriving a set of ‘behavioral rules’ (or, in some cases, having
those built in). This will enable behavioral prediction when
the same or a highly similar situation arises again. But
anotherway to do it – indeed amore flexible way to do it – is
to discern directly what the other is trying to do, what state
of the environment he is trying to bring about and what his
goal is. This enables behavioral prediction not only in
previously observed or highly similar situations but also
panzees’ understanding of visual perception, especially
from the well-known studies of Povinelli and Eddy [6] in
tive evidence also accrued during the 1990s about chim- group, highly social animals, such as chimpanzees,must be
Does the chimpanz
mind? 30 years late
Josep Call and Michael Tomasello
Max Planck Institute for Evolutionary Anthropology, Deutscher
On the 30th anniversary of Premack and Woodruff’s
seminal paper asking whether chimpanzees have a
theory of mind, we review recent evidence that suggests
in many respects they do, whereas in other respects they
might not. Specifically, there is solid evidence from
several different experimental paradigms that chimpan-
zees understand the goals and intentions of others, as
well as the perception and knowledge of others. Never-
theless, despite several seemingly valid attempts, there
is currently no evidence that chimpanzees understand
false beliefs. Our conclusion for the moment is, thus, that
chimpanzees understand others in terms of a percep-
tion–goal psychology, as opposed to a full-fledged,
human-like belief–desire psychology.
Introduction
In 1978 Premack and Woodruff asked, ‘Does the chimpan-
zee have a theory of mind?’ In this brief review we attempt
to answer this question based on much research that has
been conducted in the 30 years since that time, particularly
in the last decade or so (see Tomasello and Call [1] for the
state of the art as of the mid-1990s). The answer will not be
a simple yes or no, however, because part of the progress
that has been made in recent years is a recognition that
there are many different ways in which organisms might
understand the psychological functioning of others.
A brief history
Premack and Woodruff’s [2] original study was actually
about chimpanzees’ understanding of human goals. But
soon therewas new research suggesting that perhaps these
results were experimental artifacts [3,4] and other
research on social learning that suggested chimpanzees
did not have an understanding of human goals [5]. Nega-
Corresponding author: Call, J. (call@eva.mpg.de).
1364-6613/$ – see front matter � 2008 Elsevier Ltd. All rights reserved. doi:10.1016/j.tics.2008.0
e have a theory of
tz 6, D-04103 Leipzig, Germany
go beneath the surface to an understanding of the goals,
perceptions, knowledge and beliefs that guide action (nor
to an understanding of underlying physical forces either).
But not all of the data existing at that time were consistent
in suggesting this conclusion, and moreover, chimpanzees
had been observed doing things that would seem to require
more than just an understanding of surface-level behavior,
for example, tactical deception [9].
The story since the late 1990s has been one of exper-
imenters finding better ways to tap into what chimpanzees
know about the psychological states of others – and so
getting many more positive results. In most cases this has
been guided by attempts to model the experiments more
closely on situations that chimpanzees routinely encounter
in their natural environments, for example, presenting
them with problems not in situations in which they must
cooperate with others but, rather, in situations in which
they must compete with others (see Box 1). Skeptics still
abound, as represented most prominently by Povinelli and
colleagues [10,11]. They cling to the hypothesis that chim-
panzees understand only surface-level behavior (forming
‘behavioral rules’), and indeed this explanation is almost
always possible for any single experiment. But there are
now in many cases multiple experimental paradigms all
aimed at a single psychological state – each presenting
chimpanzees with a highly novel problem – that makes the
positing of learned behavioral rules a difficult explanatory
strategy [12]. Here, we review current experimental evi-
dence about chimpanzees’ understanding of: (i) the goals
and intentions of others and (ii) the perception, knowledge
and beliefs of others.
Understanding goals and intentions
To compete and cooperate effectively with others in their
in novel situations.
The methodological problem is that when an actor acts
toward a goal successfully, then what he is trying to do and
2.010 187
Box 1. Assessing visual and auditory perspective taking in
chimpanzees
Hare et al. [26] had chimpanzees compete for food with a human
competitor who was inside a glass booth (see Figure I). In the main
study chimpanzees preferentially chose to approach the food that
was on the side of the booth with an opaque barrier so that the
human could not see them approaching. They, thus, attempted to
influence what the human could see, and they did so from the very
Review
what he actually does are the same, and so it is very
difficult, if not impossible, to distinguish when an observer
is reading the actor’s behavior versus reading his goals.
But there are situations in which what an actor does does
not match his goal, specifically, in unsuccessful attempts
and accidents. The best evidence that an observer under-
stands goals, therefore, is when she reacts specifically to
the actor’s goal and not to his overt behavior when he is
trying unsuccessfully or having an accident.
first trials. Melis et al. [27] went one step further by having a booth
with opaque barriers on both sides, but chimpanzees had to choose
between reaching through a clear tunnel, in which case the human
competitor could see their reaching arm, and an opaque tunnel, in
which case she could not. Even though they could not see the
human under any conditions, the chimpanzees reached more
through the opaque tunnel than the clear tunnel – presumably
imagining the perspective of the human. In an extension of this,
Melis et al. [27] used this same booth arrangement but with two
clear tunnels leading to food. Here the experimenter lowered her
head between her knees so that she was unable to see the
chimpanzee. In this case each tunnel had a small door midway
through, one of which was very noisy and one of which was silent
(they had learned about the doors’ properties earlier in a different
context). If the human heard a door opening, she would raise her
head and grab the food before the chimpanzee had a chance, but
not if she could not hear a door opening. Chimpanzees in this case
preferentially reached through the silent door, and again they did so
from the very first trials.
Figure I. Experimental setup for the studies of Hare et al. [26] and Melis et al.
[27].
188
Box 2. Chimpanzees infer a human’s intentions
Buttelmann et al. [28] used the Gergely et al. [29] method to test six
human-raised chimpanzees in the so-called rational-imitation para-
digm. The chimpanzees were shown how to operate an apparatus to
produce an interesting result (e.g. lights or sounds), and then they
were given a turn. The most natural behavior for them in all cases
was to operate it with their hands. But this obvious behavior was
never demonstrated for them; they always saw a human manipulate
the apparatus in a novel way with some other body part. The idea
was that in some cases the physical constraints of the situation
dictated that the human (referred to as ‘E’ in the figure) had to use
that unusual body part; for example, he had to turn on a light with
his head because his hands were occupied holding a blanket or he
had to operate a light with his foot because his hands were occupied
with a heavy bucket (see Figure I). When the chimpanzees saw this
forced use of the unusual body part, they mostly discounted it and
used their hands as they normally would (because the constraints
were not present for them). However, when they saw the human use
the unusual body part when there was no physical constraint
dictating this, they quite often copied the unusual behavioral means
themselves. If we interpret this experiment the way it is interpreted
Trends in Cognitive Sciences Vol.12 No.5
Table 1 lists ten separate studies suggesting that
chimpanzees understand others’ goals. At least six of these
aremodeled closely on studies with human infants (see Box
2), which are routinely used as evidence of goal under-
standing. These ten studies involve several different situ-
ations that chimpanzees must understand and several
different response measures as well. Thus, in the first
seven studies in Table 1 (briefly described there, points
1–7), chimpanzees react not toward the actual behavior of
the human but toward his goal, as they attempt to get a
reward or anticipate his behavior. For example, they react
differently to very similar behaviors when the human is
refraining from giving food because he is unwilling versus
unable to do so (Table 1, points 1 and 3), when he is doing
something on purpose versus by accident (Table 1, point 2)
or else when he is an innocent bystander (Table 1, point 7).
They also discern that another’s goal is to reach an out-of-
reach object and then they either help him to reach it
(Table 1, point 4) or grab it first if it indicates the location
of food (Table 1, point 5). And they anticipate a human’s
for human infants, the conclusion is that the chimpanzees under-
stood not only what the experimenter was trying to do (his goal) but
also why he was doing it in the way he was doing it – the rationality
behind the choice of the plan of action toward the goal. According to
Tomasello et al. [30], an understanding of the action plan chosen
toward a goal constitutes an understanding of the intention.
Figure I. Turning on the light with the foot because (a) E wanted to or (b) E had
to (because his hands were occupied) in the Buttelmann et al. [28] experiments.
nd
op
imi
bo
ive
stan
impending actions when he stands and turns in a certain
direction based on what goal he is probably pursuing
(Table 1, point 6). Importantly, in two of these studies
(Table 1, points 3 and 6) the behavior at the time the
chimpanzee must react is identical in experimental and
control conditions, with the only difference being in the
immediately preceding context (e.g. given that the human
has been opening boxes, the current ambiguous behavior is
probably an attempt to open a new box, whereas without
this preceding context the human’s goal is unknown).
One could of course attempt to explain any one of these
results in terms of behavioral rules that chimpanzees are
either born with or learn, and this might be plausible in a
particular case. But this explanatory strategy is not plaus-
ible across all of the seven studies because of the diversity
of situations and reactions required. Moreover, to explain
the studies in which the human’s behavior is identical in
experimental and control conditions at the time of reaction,
one would need to also posit something like ‘contextual
rules’ because there is no differential behavior to read in
the two situations.
Table 1. Studies on chimpanzees’ and human infants’ understa
Studies
Getting/finding food
1. Leave earlier and beg more intensely from an E who is unwilling as
(behavior similar in the two cases)
2. Select the box acted on intentionally versus accidentally (behavior s
3. Leave earlier when E is playing with as opposed to trying to open a
two cases)
Reacting to a partner’s actions
4. Give the object that the E is trying to reach
5. Take the food that a competitor is trying to reach
6. Anticipate where E is going based on potential goals available
7. When food is stolen retaliate against thief, not against innocent rece
Imitation
8. Produce target action based on observing a failed attempt
9. Copy intentional actions more often than accidental actions
10. Selectively copy freely chosen acts but not those forced by circum
a(D. Buttelmann et al., personal communication).
Gaps in the table indicate no information available.
Review
In three further studies in Table 1 (points 8–10) expla-
nations in terms of behavioral or contextual rules are not
possible. This is because these all use an imitation para-
digm in which the chimpanzee subject actually acts out in
her own behavior what she understands the other to be
attempting to do, and this does not always correspond to
the overt actions (note that all three of these studies use
human-raised chimpanzees, who are very probably the
only ones capable of copying actions in specific ways
[13]). In these three studies (all modeled on similar studies
with human infants), chimpanzees imitated what the
human was trying to do (not what he did) (Table 1, point
8), his purposeful rather than his accidental actions
(Table 1, point 9), and they even selectively imitated
actions based on an understanding of why the actor chose
this particular action (Table 1, point 10), which might be
construed as understanding his intention, in the sense of
the action plan he chose for pursuing his goal. Behavioral
and contextual rules only help a subject to predict what an
actor will do; they provide no guidance whatsoever for
social learning.
We believe that there is only one reasonable conclusion
to be drawn from the ten studies reviewed here: chimpan-
zees, like humans, understand the actions of others not just
in terms of surface behaviors but also in terms of the
underlying goals, and possibly intentions, involved. Beha-
vioral or contextual rules might be concocted to explain the
results of any one of the seven studies in which the chim-
panzees react to or predict the behavior of others, but this
requires many different ad hoc behavioral and contextual
rules for which there is absolutely no positive evidence.
Indeed consistent use of this explanatory strategy would
also deny human children an understanding of goals and
intentions because most of the chimpanzee studies are
modeled on child studies. Moreover, the three imitation
studies would not seem to be amenable to behavioral-rules
explanations at all.
Understanding perception and knowledge
To understand how another works as a goal-directed agent,
an observermust understand not only his goals but also his
perceptions because what he sees and knows helps to
ing of goals and intentions
References
Chimpanzees Infants
posed to unable to deliver food [31] [32]
lar in the two cases) [33] [33]
x with food (behavior identical in the a
[34,35] [34]
[36]
a
r of stolen food [37]
[38,39] [40]
[38] [41]
ces [28] [29]
Trends in Cognitive Sciences Vol.12 No.5
determine what he does. Here, we examine what chimpan-
zees understand about what an actor sees – not just what
he is oriented to, but what he registers from the environ-
ment in ways that affect his actions – and about what that
actor knows in the sense of information he has previously
registered that still affects his current actions (e.g. his
knowledge of where food is even though he cannot see it
now). Understanding false beliefs is the special case in
which an observer predicts or explains the behavior of an
actor based on a judgment of what that actor believes to be
the case, not what really is the case as the observer knows
it (e.g. the actor believes the food is in one place when the
observer knows that it is really in another).
Again in the current case, the main alternative hypoth-
esis that we must consider is that chimpanzees either are
born with or learn certain behavioral or contextual rules
that determine how they respond to others’ surface beha-
viors (orienting behaviors in this case) without any un-
derstanding of their perception or knowledge. And
analogous to the case with goals, the best evidence that
an individual understands another’s perception is when
189
nd
on
indo
hem
ctio
f b
ood
the two individuals’ perceptions differ (e.g. they see
different sides of a barrier), and the observer acts on the
basis not just of what she herself sees or knows but also on
the basis of what the other sees or knows.
Table 2 lists 16 different studies relevant to the question
of whether chimpanzees understand what others see and
know. The first set of studies (Table 2, points 1–7) involve
gaze-following behaviors, which are, admittedly, not such
powerful evidence of an understanding of perception. The
key fact here is that chimpanzees follow the gaze direction
of others behind themselves, around barriers and past
distractors; they check back with the gazer if nothing
interesting is to be found in the indicated direction, and
Table 2. Studies on chimpanzees’ and human infants’ understa
Studies
Gaze following
1. Follow gaze to distant locations behind self
2. Follow gaze on the basis of both face and eye direction
3. Check back with gazer if nothing relevant at the target location
4. Stop looking after a few trials if nothing relevant at the target locati
5. Ignore distracting objects on the way to the target location
6. Move to the side of opaque barriers to view the target location
7. Understand that gaze stops at an opaque barrier - unless it has a w
Gestural communication
8. Use visual gestures mostly when conspecifics or E are oriented to t
9. Position oneself to gesture in front of others
10. Both face and eye orientation of recipient determine gesture produ
Food competition
11. Pick the food that the E is not looking at
12. Pick the food that a dominant individual or E cannot see because o
13. Visually conceal approach to food (using barrier)
14. Auditorially conceal approach to food (choosing silent door)
15. Take food that a dominant individual did not see being hidden
16. Understand that if competitor picks first, he will have chosen the f
being hidden
a(J. Kaminski et al., personal communication).
Gaps in the table indicate no information available.
Review
they eventually stop following if an individual always
gazes at nothing interesting. These are all characteristics
of gaze following shared with human infants, and even
though human infants use the eyes, as opposed to the head,
more often than chimpanzees do in following gaze direc-
tion, chimpanzees do use the eyes to some extent as well
(Table 2, point 5).
The second set of studies involves gestural communi-
cation. The general finding is that chimpanzees take into
account the visual orientation of the recipient when ges-
turing (Table 2, point 8), even moving themselves in front
of the recipient to gesture when necessary (Table 2, point
9). Of particular importance, Kaminski et al. [14] found
chimpanzees muchmore sensitive to the recipient than did
Povinelli and Eddy [6] in a similar paradigm when the
human was actually in a position to deliver food (i.e. facing
them bodily). It is true that chimpanzees are not as sensi-
tive to the eyes as are human infants, but still they under-
stand when someone is or is not in a position to receive
their communicative act (Table 2, point 10) – again in a
manner very similar to human infants.
By far the most power
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