Mental states in animals: cognitive ethology

Mental states in animals: cognitive ethology Jacques

Vauclair

This artHe

addresses

ethology”.

the quegtion

In effect,

behaviours

of mentaJ states

thk field of research

such as food caching,

communication

in wild

self-aw&&reness

and intentional@.

Cognitive

Mentional

occurring

imitation,

tool use and

of mental

experiences,

use some philosophical

ethologists

Moreover,

between cognitive

of cognitive

animal psychology)

ethology.

in ‘cognitive

stance’) to carry out their programme of the

other approaches to the investigation of cognitive

naturally

recognition,

of natural behaviours. A comparison

(e.g., experimental

as viewed

in order to seek for evidence

cencepts (e.g., the ‘intentional investigation

aims at studying

individual

animals,

in animak

ethology and

processes in animals

helps to point out the strengths and weaknesses

laboratory

attempts to analyse experimentally

behaviours such as deception, the relationship

between seeing and

knowing, as well as the ability of animals to monitor their own states of knowing, suggest that cognitive frameworks

ethology could benefit significantly

and methods of animal cognitive

fact, co&ribute

to the understanding

from the conceptual

psychology.

of which cognitive

Both disciplines could, in abilities are evolutionary

adaptations.

T

he term

Griffin

‘cognirive

veloped

in other

book

a strong

an extension relying

Frisch’,

chimpanzees”,

Griffin

interspecific

on to animal Briefly, tionary

concluded

environment

with

study

these

claims

in

beliefs.

information

in animals-.

Cognitive

to express

themselves study

concerns in canids.

intraspecific

traditional

belirz~r

that Y should

pertain

exchanges,

by cognitive namely

is related

already

However.

CopyrIght

different non-human follow

firsr-

Alarm

to the situhim: X

finally,

‘second-

wantsY

of CE is

the piping

X UUMS Y to of bow

behaviours

or perhaps

second-order

calls of vervet

monkeys

have similarly

to

been used to

and level of communication evaluation

in these of the use

stance in CE. see the commentaries

Ltd All rights Cogmtlve

reports

characreristic

plover

(Charadviw

reserved

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00 Vol

that

of a shorebird,

when

confronted

PII: Sl364.6613(97)01004-8 1.

No

to

‘third-order’

in which

(for a critical

for

a bow

Ref. 13).

by

I”

express

primates”

of the intentional Ristau’

Trends

applies

to play with

to a case where

classes of predators”

assess the complexity

Science

describes,

dog X performs

the situation

behaviour.

the play

0 1997. Elsevier

intentionality

him:

ate

has been

suggests that play-soliciting

dogs might

to inter-

investigated

the novelty

BekofP”

intentional

here. The

categorization

play with

qualifies

(see Fig. I);

that X wants Y to play. Based on the analysis

behaviours.

antipredator strategies in birds. the cognitive ethological ap-

to activities

ethologists;

Mew

the underlying

levels of intentionality

wantsY

applies

stance’)“.

As an illustration, animals”

intentionality

dog X

pro-

intentionality

several

in which

‘first-order’

intentionality

have,

between

: ‘0 order’

order’

in the natural

briefly

Bekoff”

in which

among

and the second topic

specific exchanges, namely Behaviours analysed by

ation

analyses.

following

the situation

cog-

and. consequently,

Two of them will be presented

often

abilities

have thus been investigated

intention

from

frameworks

a scale for evaluating

The

genetically

behaviour?.

USC conceptual

exchanges

postulated.

thus

hexible

(such as rhe ‘intentional

cognitive

ofsocial

example,

the evoluammal

their

has offered

drawn

on field studies.

topics

realizing

interpretation

of some fixed

more

ethologists

in the case of social play,

a ‘window’

about

with

by philosophers

Bennett

intraspecific

of non-human

of the species under

first topic

more

complexity

or Intentional

are a mixture

elements

movement:

chance

CE relies mainly

proaches

by

described

constituted

consciousness.

a better

ethologists.

for

minds. CE is concerned

cessing and rationality

Several

as

communication

that

communicarions

processes,

apviori,

gestural

provided

goal ofexplorThus.

a purposive

which

(Cognitive in

perceived

of animals.

it advances

for activities

reacrion

such as bees’ ‘language’.

and comparative

nitive

was essentially

that

transmitred

behaviourism

the ambitious

or trained

by IS’6

salutary)

by strict

experiences

on achievements

Von

and

with

mental

Griffin‘s

certainly

the book

of ethology.

ing the conscious

‘_ Although

imposed

of animals,

was comed

Au~aarmess’ and later de-

(and

the inhibitions

the study

(CE)

ofAnimd

publications’

was first

against

by

ethology’

in The Question

1,

April

1997

cd:+33491 1642 73 fax+334917149 38 e-mail: vauclai~lnf. cnrs-mrs.fr

Vaucla~r

Mental

states

in

anlmals

ceptualized

without

representation such

reference

concepts

have

and heuristic

barrier in

this

the position

A young

in baboons.

of ‘second-order’

baboon

(A)

she eats rhizomes that loudly. thus attracting

A is looking his mother’s

chases

young

T away

and

food.

The

(who

has previously

the

interpretation enacted

looks

is not

at a female

the this

study

(T) while

for. Suddenly, A screams attention: the mother B

baboon

of

intentionality

has free

access

young

baboon’s

sequence

with

that

individ-

he made his mother believe that the female to him, thus leading her to intervene and

enable

for the desired

him to reach from

food

(drawn

even

itself

parative

that

intruders

perceived

tion displays away

as potential

have been documented

from

either

of such protective

display’.

This behaviour

from

a fanning

arched

wings

her analysis

the behaviour

plovers’

distraction

nest or from

nature

forms

displays

intentional

the plover,

claims”

behaviour) analysis)

terms, according by performing away

of

to the

ual recognition,

behaviours

provides which

imitation.

presume

that such a subjective

it would

the existence

but rather

of mental

of the

food-caching.

individ-

indices

tool use and communication”‘.

ethology

and the comparative

study

It must

be above

that

said

the definition

is extensive

search areas, such as comparative tive cognition

commit

to identifying

rwo

to encompass psychology

re-

and compara-

lated”

this was first shown that self-recognition

the ability others.

their

Self-recognition

might,

about

ity of the animal

to represent

processes,

to

ity to use novel,

displaced

name a few, are at the heart of the experimental

approach

of

state and behaviour’“.

As alternative for mirror

comparative

cognition

in general

objectives,

vented

a collaboration

the cognitive that

Trends

animal

the high

in

reliance

Cognitive

Ii Notwithstanding

the convergence

could

possibly

two

have so far pre-

other

animals),

main

obstacles

between

the cognitive

psychologist.

First,

of CE on thought

Sciences

-

Vol

ethologist

and

it is very surprising processes

1.

No.

is con-

1,

April

concluding

account further

1997

on animal

its own body,

in a specuand

states of

or else the abilabout

its physical

theoretical recognition

demonstration

hypotheses in apes (and

is required

have human-like

experiences.The thinking

later

the mental

visual feedback

that these animals

ness and mental position

who

in fact, only express an abil-

of information

solving

see ref-

own reflections

lead to self-awareness

could

inferences

fallacy

self-recognition.

by GallupL”

to make

be

CE and its

the first

with

of

it could

and chimpanzees;

Ref. 23) recognize

the topics

of problem

In effect,

pre-

other

gorillas

of pro-

‘the ambasthe validity

sophisms:

self-awareness

and The

imprecision

to question

and thought

processing,

at large”.

and scope of CE

enough

to study

The conceptual

experiences

abil-

actions’.

that

mirror;

sented

adaptive

of their actions

a matter of mental representation”“. For example, UllmanL? has suggested

erences within

cognition

for are:

when

being

of

ethologists

of their

strategy

in animals.

might

mental

has the disadvantage

research

Ape species (orangutans, Cognitive

how

to animals,

proponents

of the

in animals

(2) their

result

in CE has, thus, led some scholars only

activities

actions;

sadors of thinking’

research

study

to clarify

in the sequences

the anticipation

mental

for us, to

it is like to be a bat.

aim of a scientific

of their

no specific

qual-

is not to find out what it is like to be

its anti-

are potential

be difficult

what

of the

at least a system

observable behaviours”‘. the criteria retained by cognitive

of such diverse

a list of some of these naturally comprise

In fact, com-

type of animal,

choice

agree

presuppose quality

animals

viding

ethologists

psychologists

investigation.

from

the

and in sharp

representations

to envision

the complexity

no reference

it is not the subjective

the principal

of other

attributing

relevant

behaviours

(I) (4)

the young.

All ecologically

Nagel”‘,

asserting

for the

requires

is under

ities; (3) the flexibility

are used are indicative

(first-order

understanding,

states cause Among

From

Ristau

to the intruder’s

of mutual

a certain

ranging

to con-

to assess. In effect, without

paraphrase minds

appeal

by some cognitive

that

writes: human

the rationale

comparative

psychologists

Fortunately,

im-

and outstretched

aims to lead the intruder

areas for CE. Bekoff occurring

walk

of these birds. In other

ethologist,

most

is the ‘broken-wing

behaviours,

(adiustments

activity,

The

and drag along the ground.

of piping

of the purposive

predator

behaviours

tail to an awkward

intense

cognitive

birds.

can take progressive

that flutter

that rhe conditions

several distrac-

to lure such predators

the nest or young

pressive

in which

predators:

Terrace

As a consequence,

if one accepts

in the cognitive

cognitive

that

reports,

Griffin’),

ity is impossible wirh

arguments

taken

experience,

contrast

the

Thus,

for using

processes in animals

notably

experience

by B.L Deputte;

upon

consciousness”“.

conscious

Ref. 45).

rationale

to the position

(most

behawour

uals) would be that had been aggressive adapted

to animal

to the

other

of cognitive

mental

at the core of the

that

are unconscious.

based

in CE’“,

with

In strong

assume

or to introspective

contrast

synonymous

in CE, most psychologists

cognition

‘Just as the modern

ethologists’.

meaning

it is nevertheless

adopted

animal

sciousness

wild

role assigned

cognitive

that have to be elucidated.

of

usefulness

of both

the central

has no precise as being

processes of animals

example

their study

by some

or mind,

phenomena

Fig. 1 A possible

comes from

concept

state, thinking

terms

demonstrated

CE

it is envisioned’

field

such as

animals”~‘“~‘V.

The second

with

concepts,

see Ref. 7), even though

in the comparative

to consciousness since

(but

amply

value

and laboratory

Although

to cognitive

or memory”

second

is related

sophism

before

self-awarein CE’s

to the idea that they

plan future

activities

that anticipatory behaviours, tional

with

detection

of regularities

that

author a kind

always

location),

and flew

might

could

topic

within

of research

investigators

impute both

a working

human and

desires

Such

for carrying

Premack’“,

its consequences

operational

framework,

conducted

experiments

can both

reacted

mented’“, know

ing about

and an agenda

specifically

among

issue. Thus,

authors

for Woodruff the

With

such an

for

example,

in chimpanzees involving

(I)an manipu-

observables

states (the ‘RAM

sug-

for deception

in the

states or whether in behaviou?“.

with

are acting

(the ‘RO

ences is then

is to

on the basis of reasonor mental

As a step towards

choos-

certain

an endeavour

control

that

in apes (or

their

edge formation.

to the presence absence

that manipulate between

visual

Such experiments

and rhesus monkeys can understand of others.

the seeing-knowing with

chimpanzees

are proof

and knowl-

have assessed in chim-

(Macaca

mulatta)”

the results

unlike

at-

the understanding perception

the consequences

Briefly,

chimpanzees,

to their

to the

own states

for subjective

ongoing

some

relationship. ” indicate

that

” how an of the visual

obtained

rhesus

indicate

monkeys, Recent

states) has recently the context

progressively

two

primary differ-

so that

identifications

limits.

response.

This

permits

response

the subject

the subject

the uncertain the subject

to escape into

delaying

ward).

would

discrimination

rhesus

an error

monkeys

in

an ability

actions examine

and

in other

the most to

demonstrated

and metacognitive Further

of their

animals

Both

that was identical

monkeys

uncertaintyi’.

if the monitoring

present

and

task’..

in the same experimental

for self-knowledge

to subjective

dolphins tasks””

escape from

an ability tested

try

uncertainty

discrimination discrimination

dolphins

the capacity

The

for testing

to sparingly

participants

Therefore,

both

a visual

trials near threshold,

setup.

to occur.

in acoustical

a food reinitially

and then the escape re-

used successfully

tmncam)

difficult

response was likely

has been

(Tursiopi

that subjects

is given response).

an easier trial but at a cost (for example. It is thus expected

is

of the stimulus,

At this point,

a way to escape (a third uncerrain

with

moni-

size of the stimulus

reduced,

difficult

close to perceptual

of uncertaintv

is presented The

been inves-

own

studies behaviours

species (comparable

reshould is also

abilities

might

also exist in rats) “I. Such directions

of research

should

mately

to establish

the relation-

help cognitive

psychologists

ships between

body

and awareness

of others’

self-awareness,

cognitive

mental

ulti-

self-awareness

states.

is not

of mental

in monkeys)

within

with

Concluding related

connections

tigations

com-

of research

interpretation’)

be performed,

by experiments

understand

they have

undertaken.

vided

some

have

As Heyes

this kind

interpretation’).

causal

that

and respond to check

that of human

For

the chimpanzees

must

experiences

they demon-

in the chimpanzee.

experiments

panzees”

on a num-

whether

whether

suggestions

animals

to the mental

can also be directed

monitor

paradigm

false information,

one of these interpretations,

systematically

attention

is, subjective

whether

respond

capacity

This

sponse when

chimpanzees

have been criticized

the animals

explores

of knowing.

the primary

and a human

the trained

a real capacity

to differences

of mind

appropriately

ways that animals

confronted

when

have,

a chimpanzee that

non-

of the message and

ing between

individual

and

the subject

sets of mental

whether

tributions

the theory

responses.

setup.

the basic problem

Other

Whereas

discrimination

for true deception

merely

of

of experiment,

strate

attributed

the

studied

and it is questionable

really

for attention

or to con-

on the receiver.

one may wonder

the subjective

gaze-

mind.

species showed

an ability

of the difficulty

to represent

be-

the

to propose

ber of grounds, example,

differences

(as seen in

to himself

can be inferred

experiments

possible

has led scientists

chimpanzee. The above

and apes

the perception-

“. Uncertainty can, for example. be experimentally in a discrimination problem. Typically. in this type

and produce

to the authors,

monkeys

in animals

between

understand

world.

toring“’

of mind’.

the content

show

experi-

tigated

this

with

that an individual

on deception

Results

in

(2)

because

primate

the other

between

chimpanzees

experiments)

the the

to at least TWO discrep-

to understand and

and

to

can

these

of information

following non-human

point

differences

states of conspecifics,

dominant

particularly

findings

relationship.

humans

eyes connect attention

In this

in a communicative

experimenter.

knowledge

the of

ability

of a

(that

intentionality

understands

respective

in the orientation

(see also Ref. 35). do not

cognition

(more

to address

(1) possible

can understand

a controversial

for intentionality

sender of a message controls

gesting

is, indeed,

or beliefs

experiments

primates)

lation

intention.

‘the theory

an inquiry definition

out

troglodytes),

to anticipate

the above

ancies:

tween

how states

in their

where

the ability

search for evidence

intentions,

specific?.

the

the last visited

psychology,

into

Altogether,

Some bees

location

this issue has also become cognitive

It is

and conscious

of inrentionality

animal

the advent field,

again,

internal

that the ex-

beyond to a new

planning

to appreciate

a food

to shifts

by a experimenter

appear

to trial:

behaviour.

the food

Here

with

CE. Interestingly,

trial

(as if they understood

immediately

be found.

The question

from

can respond

menter’s

in his studies

bees to fly towards

they

the

and anticipate

Frischj

of anticipatory

placed

be confused

detect

by Von

its location

these changes

perimenter food

quickly

events.

although

gaze performed

of inten-

begins with

environmental

trained

changed

bees displayed anticipated

the intervention

as observed

This

which

will

It appears

of most adaptive

as adaptation

among

an organism

on insects. source

imply

processes,

these regularities,

intentions”.

are at the heart

and do not

or conscious

likely

conscious

activities

do inves-

these primates.

Trends

remarks

CE has undoubtedly exploring their

‘mental

lacks research fulfil

contributed

the complexity experiences’.

However,

and experimental

programme.

exist and they derive

from

and

concepts

success by those biologists

or ecological

approach of cognitive

I”

Cognltlve

cognitive

of the psychological

cessful

Scfencer

field

currently

approach

to animal

-

Vol.

and

methods

can be used in the synthetic

behaviour”‘.“.

mechanisms

to

actually

psychology Both

interested

in and

paradigms

of behaviour”.

with

study

this

Yet, these methods

human

analysis

of interest

life of animals

methodology

its ambitious

the experimental

to the wave

of the social

The

involving

1, No.

1,

suc-

naturally

April

1997

Table

1. Cognitive

ethology

Cognitive

versus

comparative

ethology

The study of natural behaviours settings from an evolutionary perspective

Scope

To investigate minds, processes, rationality

Methods

The use of anectodes and empirical with ethological techniques with different species issue

imporranr

social

play”).

According about

using

logical

memory

that

us to reconstruct

significant

hehaviou?

and

our

psychoethol-

1). It can be ex-

comparative

approaches

own

opposition

about

cagnirive

experiments.

The

will

features

experiments Second,

tory

as nored to provide that

nipulating

experimental

’ I-. Two points

phenomena solutions

on the basis of a minimal

obser-

in free-living

The merge

within

including

intenthe latter,

psychology

experiments believe

and

in

cognition

in the debate. manifests

problems”“.

First,

cognitive

processes

in an ethological

on animals

and

Finally, animal

in

Cognitive

Sciences

Vol

1,

No.

1,

April

In particular,

as

the study

a cog-

introducing

component

of

research

between

into

that

etholoo

raises the importanr

animal

rights.

Inevitably.

wherher

animals

feel, think

and

quests to determine experiences

fuel the debate

of the time these questions

nificant

to animal example

given

full

those

scientists

rights

where

human

humans

is truly

who

have rarely

entered

study

over animal

the cognitive

the debate

their data and interpretarions

and have

righrs

that apes should astonishing capacities

or cognitive

be that

of non-

psychologists)

(but see Refs 43,50), would

issues.

by activists

(see Ref. 48 for a sig-

propose

Ir is, however,

ethologists

issue of repeated

are addressed

movements

activists

rights).

(cognitive

where

be very helpful. I..

References 1 Griffin,

D.R. (1976)

University 2 Griffin,

1997

The Question

of Animal

Awareness.

Rockefeller

Press D.R. (1978)

1, 527-538

Trends

might

and role of

through

thus

an ecological

cognitive

welfare

belonging

CE could help place the analysis of cognition within an evolutionary framework51,52 and thereby help us to understand the biological framework in which cognitive processes have evolved. l Cognitive ethologists claim’that the consideration of individual behaviour is often neglected in favour of the study of the behaviour of the species. The study of a few individuals could verify the existence of strong intraspecific variations and reduce generalizations about the cognitive skills of the whole species. *The expression of cognitive abilities by animals in their environment and their sensory world needs to be investigated. This growing field of ‘sensory ecology’53 should aim at ‘taking the animals’ point of view’ by studying their information processing in conditions as close as possible to those in which they live. l

studies

perspective,

and its environment,

science

of

needed.

Most

questions

er al.‘“,

may shed lighr on the coupling

system

mental

Outstanding

sciences.

ma-

in successful

Use of anecdotes

cognitive

and

into the laboratory itself

and field

Previde

social

all, behaviours

the importance

by Prato

nitive

we have

states are formed

(after

proposed

laboratory

is not inconsistenr

also have a history).

laboratory

animal

easily be transferred

unusual

between

at work,

variables,

often

experience

conditions

The arguenvironments

Moreover,

rhat mental

in recognizing

“‘.SF, whereas

in artificial of training

so far, to prove

of

are less

states in the labora-

of intentionality.

help significantly

control

can be raised

intelligence to

that field

of intentional

appearing

the attrihution

animals

the credo animals

needs to be demonstrated.

hehaviours

cognitive

cannot

social

claim

of experimental

of performing

ethologists

of field versus

help design

in the laboratory.

to which

are the result of a long history with

a first step to

could

and Dickinson’.,

according

evidence

that

of variables

by Heyes

ethologists,

than in the field.

ment

thus constitute events

for later conrrol

cognitive likely

might social

to see the true complexity

of belie&

the tradition

the necessity

virtues

former

of social exchanges

and attributions

in line with

between

the respective

offer the best opportunity

and intricacies

and

(see Table

how

is a longstanding

laboratory

work’

observations

document

and methods.

processing in and non-verbal

To test Darwin’s gradualist hypothesis concerning the continuity between human minds and the minds of other animals

qualitative

ir is now

of

Use of the methods and paradigms of experimental psychology (e.g., reaction times), mostly in laboratory settings

no evidence.

and psychologists

tionaliry

gained

(for

integrarive

and continuity species

To investigate information memory, problem solving thought

song

including

theories

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Perceptions

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Trends

I”

Cognitive

Sciences

Vol

1,

No.

1.

April

1997