Charles Whitehead

The Neural Correlates of Workand Play

What Brain Imaging Research and Animal

Cartoons can tell us about Social Displays,

Self-Consciousness, and the Evolution of the

Human Brain

Abstract: Children seem to have a profound implicit knowledge of

human behaviour, because they laugh at Bugs Bunny cartoons where

much of the humour depends on animals behaving like humans and

our intuitive recognition that this is absurd. Scientists, on the other

hand, have problems defining what this ‘human difference’ is. I sug-

gest these problems are of cultural origin. For example, the industrial

revolution and the protestant work ethic have created a world in

which work is valued over play, object intelligence over social intelli-

gence, and science and technology over the arts. This may explain

why we have so many imaging studies of tool-use and object manipu-

lation, but only four studies of dance, two of pretend play, and one of

role-play.

Yet in order to understand child development, the evolution of the

brain, and the emergence of human self-consciousness, we need to

look at social displays — such as dance, song, image-making and

role-play — which underpin human culture, cooperation and the arts.

I will discuss recent brain imaging research on playful versus instru-

mental behaviour and show how, in conjunction with archaeological

data, we can use this to make sense of human evolution.

Journal of Consciousness Studies, 15, No. 10–11, 2008, pp. ??–??

Correspondence:drcwhitehead@aol.com

Everyday Pretence

We humans are much concerned with managing our self-image in

order to influence other people (Goffman, 1959) — so what we call

personality and selfhood reflects a kind of ‘marketing and branding’

campaign. This everyday self-promotion depends very much on pre-

tence and make-believe (Turner, 1982). Most children start to engage

in pretend play somewhere between the ages of nine and eighteen

months, and I believe this scaffolds the development of self/other

awareness (‘theory of mind’) and imagination (‘theatre of mind’)

(Whitehead, 2001). Pretending is something children do for the sheer

fun of it, and children who cannot pretend by the age of eighteen

months are likely to be diagnosed as autistic by the time they are four

years old (Baron-Cohen et al., 1996).

One thing no normal human being wants to be perceived as by oth-

ers is an animal. To call someone an animal or a beast is deeply insult-

ing, and this appears to be true cross-culturally (cf. Lévi-Strauss,

1955). So you could say that a major behavioural difference between

humans and chimpanzees is that chimps don’t spend their adult lives

pretending they’re not apes.

We generally try to conform to the image which we think other peo-

ple expect of us (cf. Cooley’s ‘looking-glass self’: 1902), so that our

acts of pretence tend to become standardized within a given culture.

As we deceive each other, we are of course being deceived ourselves,

and the deception of others is all the more effective for involving self-

deception (Trivers, 2000; 2007). So there will be large areas of over-

lap between self-deception, ‘collective representations’ (Durkheim,

1912; Turner and Whitehead, this volume), ‘false consciousness’

(Marx and Engels, 1846) and ‘collective deceptions’ (Knight, 1991;

this volume).

Any human institution is inevitably political. It has to maintain

its power, influence, authority and credibility. This means that

politicians, lawyers, doctors, clergymen, and even scientists — among

others — have all developed their own professional schemes of ‘col-

lective deception’.

When Children Seem to Know More than Scientists

Chuck Jones — the originator of such immortal favourites as Bugs

Bunny, Road Runner and Wiley E. Coyote — is no behavioural scien-

tist. Yet, in a sense, he knows more about human behaviour than many

scientists do. In a different sense, however, we all must know what he

2 C. WHITEHEAD

knows, otherwise we wouldn’t laugh at his cartoons — where much

of the humour depends on animals behaving like humans and our

intuitive recognition that this is absurd. Chuck Jones provides a good

illustration of the difference between implicit knowledge and scien-

tific knowledge. What may seem surprising is that, where human

behaviour is concerned, implicit knowledge appears to be more reli-

able than scientific knowledge.

A major article in Current Anthropology (Henshilwood and Marean,

2003) noted that there is no agreed body of theory that can define

‘modern human behaviour’ or specify exactly how it differs from the

behaviour of other animals. There are conceptual gulfs dividing the

major behavioural sciences, and disagreement is rife. When I first

decided to study anthropology I was surprised to discover that few

social anthropologists were interested in psychology (because Durk-

heim said that ‘social facts’ were irreducible to lower orders of expla-

nation), and the same was true of biological anthropologists (because

‘proximate mechanisms’ were ‘somebody else’s job’). Further, social

and biological anthropologists did not and could not communicate

with each other. When Leslie Aiello became head of department, she

told me that some years previously the department held a meeting of

biological and social faculty in an attempt to resolve their differences.

The only conclusion arrived at was not that the two parties favoured

different ways of answering questions about human behaviour. They

could not even agree on what a question was.

How is it possible that different groups of highly intelligent and

scientifically trained people cannot agree on any definition of what

makes humans different from animals, when my children seem to

understand this perfectly well? I say this because they laugh at Chuck

Jones cartoons which are full of sophisticated insights. I will just illus-

trate with one example (Figure 1).

THE NEURAL CORRELATES OF WORK & PLAY 3

Figure 1. Two coyotes react to the sight of a rabbit behaving like a human

being (based on an unidentified Chuck Jones cartoon. Redrawn by Jon

James).

In this clip, two coyotes are watching Bugs Bunny. He swaggers

along a road, smoking a cigar and showing off to the world his smart-

Alec superiority. His every gesture is a display of material, intellec-

tual, social and cultural wealth. The coyotes’ reaction to the sight of a

rabbit behaving like a human being is of course a double joke — they

themselves are behaving like humans. Yet how long did it take scien-

tists to explicitly acknowledge that even our closest relative, the chim-

panzee, lacks the surprised facial expression? And I am not aware that

any scientist has yet reported that chimps do not check out each

other’s faces as if to say ‘Did you see what I saw?’, yet we surely

know that they don’t.

Generally we expect scientific knowledge (e.g. the earth is round)

to be superior to intuition (e.g. the earth is flat). But we humans have

evolved very high levels of social insight and social intelligence, so

that our intuitive knowledge of human behaviour is generally reliable.

Scientific knowledge, on the other hand, is curiously blinkered. The

likeliest explanation for this disparity, I suggest, is active falsification.

Animal cartoons and worldwide ethnographic research implicate

three major behavioural differences between humans and other ani-

mals. The first is our formidable armamentarium of social displays,

ranging from the surprised faces of the cartoon coyotes to more com-

plex cultural displays such as Warner Brothers movies. The second is

our associated level of self-consciousness and insight into the con-

scious states of others — without which the surprised face and cartoon

movies would serve no function. The third is economico-moral cul-

ture — that is, all human societies are structured by formal systems of

‘exploded’ kinship (clan membership, nationality, ethnicity, etc) and

reciprocity (gift exchange, trade, marriage, sporting contests, etc).

Human cooperation depends on our readiness to identify with each

other, together with these expansive structures; and the price we pay

for the latter is a life of masquerade and collective deception (White-

head, 2002; 2003; 2006a, b).

Social Display and Self/Other-Awareness

At the consciousness conference ‘Tucson 2006’, Temple Grandin

gave a talk entitled ‘I Think in Pictures Instead of Language’.1 She

explained, for example, that if someone says ‘church steeple’, she

would understand this by running a mental slide-show of actual

church steeples that she has seen in the past. A mind that thinks in

4 C. WHITEHEAD

[1] Temple Grandin is famed for her central role in Oliver Sacks’ (1995) An Anthropologiston Mars, and as the designer of uncannily successful livestock equipment.

pictures, apparently, has no generalized abstractions, only specific

images of real-world objects and events.

Consider the implications. If, for example, I had to render the title

of this paper in pictures instead of words, I would have to skip articles,

conjunctions and prepositions. There are no syntactical relationships

in pictures. For the word ‘Neural’, I could show a drawing of a

neurone: but this would imply a noun, not an adjective. Concrete

adjectives such as ‘big’ and ‘green’ have obvious visual equivalents,

but the abstract concept ‘pertaining to’ does not. The associated noun

‘Correlates’ is another abstraction with no obvious visual equivalent.

Anyone who has played ‘charades’ will be familiar with the prob-

lems of non-verbal communication. In a game of charades, to indicate

the word ‘Work’ I would have to mime specific concrete examples

such as digging or hammering, and it might take some time before

people arrived at the generalized abstraction ‘work’. The word ‘Play’

faces the same problem. The toughest word of all is probably ‘Con-

sciousness’ — something we are all sure we possess but can never say

what it looks, sounds, feels or smells like.

Representing something by resemblance — as in drawing a picture

of a neurone, miming someone digging, or mimicking the sound of a

police siren — is known as ‘mimesis’. You could say that mimesis is

more ‘primitive’ than language. Apes use mimetic signals spon-

taneously (Tanner and Byrne, 1996), whereas they have to be taught

to use verbal signs. Although fully articulate mimetic gestures and

vocalizations appear in children around the same time as the first

words — at about twelve months — simple indexical gestures and

‘clowning, tricks and jokes’ begin six months earlier (Trevarthen,

1995). But ‘primitive’ does not always mean ‘inferior’. Mimesis cer-

tainly has serious problems if you try to force it to do the same job as

language, but that is because the two modes of communication have

different functions. If you want to demonstrate something concrete,

mimesis is generally superior to language. To tell someone how to get

from A to B, the best way is to draw a map. To explain how a petrol

engine works, or how to use a mallet and chisel, diagrams and demon-

strations are worth more than a thousand words. One reason why

PowerPoint seems to be taking over the world is because scientific

communication would be severely impaired without the use of graphs,

charts, scans and images of many kinds — all of which come under the

heading of ‘mimesis’.

Mimesis also has functions other than communication. If a child is

pretending that a banana is a telephone, then the banana is a mimetic

representation. But pretend play has no communicative goal — it is

THE NEURAL CORRELATES OF WORK & PLAY 5

autotelic: that is, self-motivated or ‘just for fun’. In contrast to the

problems encountered in games of charades, pretend action is very

easily understood by normal people. It takes the average adult less

than three seconds to understand a pretend action.2

There are two main clues that can tell you when someone is pre-

tending. The obvious one is that the object is used mimetically — the

banana is held and spoken into like a telephone, or the pen is made to

swoop and dive like an aeroplane. But you can also tell whether some-

one is being serious or ‘silly’ from their facial expression, which

infants can read from a very early age (Lillard, 2001). People gener-

ally show when they are being silly by making a ‘play face’ — with

raised eyebrows and covered teeth — which says, in effect, ‘I am not

going to bite you’. Mothers use it when they pretend to be The

Monster, and their toddlers flee squealing with terrified delight. The

same facial expression is used by chimpanzees to indicate the differ-

ence between a play fight and a real fight (Young, 1992). Mock

aggression is common among primates and social carnivores, and is

an important part of socialization. This expression of mock aggression

comes to be used to indicate all kinds of fun — such as pretending that

a banana is a telephone.

Where language scores over mimesis is in referring to intangibles —

abstract or generalized ideas and matters displaced in space or time —

that is, things imagined or imaginary. Language is a conventional

cryptic code — it refers to things, thoughts, perceptions and ideas by

consensual agreement and not by resemblance. And, whereas mimesis

is analogical — using sliding scales of infinitesimally variable grada-

tions — language is digital, requiring syntax for its infinite powers of

recursion3 (see Introduction to Knight, and Knight, this volume).

Even the sign languages used by deaf people, though many of the

words are clearly mimetic, are nevertheless cryptic and syntactical.

6 C. WHITEHEAD

[2] Based on response times of eleven adults watching video clips of three different pretendactions, prior to designing a pretend play study using similar but not the same videotapedactions (Marchant et al. in preparation).

[3] Neuronal language theorists may object to such a sharp distinction between language andmimesis (cf. Pulvermüller, 1999; Glenberg and Kaschak, 2002; Barsalou et al., 2003;Gallese and Lakoff, 2005). They may argue that if concepts are represented in sensori-motor brain areas, as are the conceptual meanings of words and sentences, then both con-ceptual thinking and language are likely to be continuous with ape thinking. This does notfollow. The conceptual abilities that enable human beings to invent and use, say, Morsecode, may well be continuous with the abilities of non-human apes, but Morse code is not.It was developed in the early 1840s for use with Samuel F.B. Morse’s electric telegraph.This historic innovation could hardly be predicted from ape abilities alone. Nor can a con-sensual system evolve by natural selection acting on traits in individuals (see Knight, thisvolume).

Modern humans use many cryptic codes other than language, includ-

ing conventional gestures both polite and rude, mathematical denota-

tions, traffic signals, and so on.

To summarize so far, I have emphasized the differences between

mimesis — indispensable for conveying specific concrete ideas —

and language — essentially committed to handling abstractions and

imaginary or imagined concepts. However, I have also mentioned a

third mode of communication. The ‘play face’ is part of our affective

gesture-call system (Burling, 1993), along with vocalizations such as

laughing and crying, facial expressions such as smiling and scowling,

and ‘body language’ generally (see Weisbuch and Ambady, this vol-

ume). Unlike mimesis and syntactical language, much of our gesture-

call signalling is involuntary, which means that it is usually honest.

Voluntary systems such as mimesis and language can be used to lie,

and could serve no useful function without the prior emergence of

sufficient levels of social trust (Knight, 1998).

So we have at least three modes of communication (Burling, 1993),

which I will call implicit, mimetic and conventional. Note that they

are associated with different levels of subjective experience: implicit

signals express affective and autonomic states and sensations; mimetic

signals convey concrete percepts and concepts; and language commu-

nicates the constructs of abstract thinking, dreaming and imagination.

Significantly, the same three modes are also found in human play and

performance.

Play is clearly different from communication. Whereas communi-

cation is goal-directed and manipulative (Krebs and Dawkins, 1984),

play is autotelic (Turner, 1982) — we engage in it purely for fun.

Performance — including song, dance, ritual, theatre and all the

cultural arts — combines aspects of both play and communication: it

is often done for fun, yet it is also goal-directed and manipulative —

aimed at an audience or at fellow performers. It also has functions

over and above those of communication and play — notably social

grooming and social entrainment (Connor, 1992; Whitehead, 2001,

2003) — encouraging self-identification with the group, and ensuring

that participants are ‘dancing to the same rhythm’ or ‘singing from the

same hymn sheet’. Performance, in effect, takes two or more ‘selfish

individuals’ and turns them into one great big selfish individual.

We humans live in a multi-layered world of shared experience,

made possible by at least three modes and three kinds of social display

(Table 1). According to social mirror theory (Dilthey, 1883–1911;

Baldwin, 1894; Mead, 1934), social displays make feelings, sensa-

tions, perceptions, thoughts and imagination salient so that, during

THE NEURAL CORRELATES OF WORK & PLAY 7

childhood development, we begin to notice that we and others have

such experiences, and so become self-conscious and other-conscious.

In other words, ‘mirrors in the mind depend on mirrors in society’

(Whitehead, 2001). According to this theory, we humans are so highly

self-conscious and other-conscious because of our formidable arma-

mentarium of social displays.

Social mirror theory has been somewhat overshadowed by alterna-

tive views. Paul Harris’s (1991) simulation theory, for example, pre-

supposes reflective consciousness as innately given, holding that we

infer subjective experience in others on the basis of our own subjec-

tive experience — self-awareness comes first, and other-awareness

derives from it. Gopnik and Meltzoff’s (1994) ‘theory theory’ invokes

simple mimicry as the basis of mindreading ability, which would seem

to make ‘mirror neurones’ and visuo-kinaesthetic matching (the abil-

ity to match a visual image of someone else’s action to a kinaesthetic

image of your own action) sufficient to explain human levels of

self- and other-awareness (cf. Meltzoff, 1990a,b; Meltzoff and

Moore, 1977; 1983; 1995).

If social mirror theory is correct, it follows that we can only be

reflectively conscious of those subjective states which are sharable

with others through some kind of display — a ‘subjective report’ in

the broadest possible sense. This may seem counter-intuitive, but it

can easily be tested. For example, why is it that the dorsal visual

stream is generally unconscious, whereas the ventral stream is con-

scious (Goodale and Milner, 1992; Milner and Goodale, 1993; 1995)?

Is it just a coincidence that dorsal vision is egocentric whereas ventral

vision perceives a universalized world — the one we share with

everyone else?4

A ‘Play and Display’ Hypothesis of Brain Evolution

It is clear that our prodigious repertoire of social displays could not

have emerged all at once, and must have evolved in a logical order.

Communication has to be the oldest type because even individual cells

exchange chemical information. Play is also fairly primitive. If you

define play broadly as ‘exploratory behaviour’ then even paramecia

exhibit this (Hameroff, 1994). If you further define play as explora-

tion of self-potential and rehearsal for adult life, then this is

8 C. WHITEHEAD

[4] The sharing of visual experience begins very early in infancy. Even newborns prefer tolook at faces with eyes directed towards them, and at three to six months will look in thedirection of another’s gaze (Csibra, 2004). Six-month-old infants will point out things thatinterest them to others using indexical gestures (Trevarthen, 1995).

widespread in social mammals and not absent from birds. Perfor-

mance is best understood as a playful extension of communication

(Jennings, 1990, 1991) so this must evolve after the other two. This

gives us an arrow of evolutionary sequence running from left to right

across Table 1. We can similarly argue that implicit displays must

evolve before mimesis, and both have to be in place before they can be

conventionalized to give rise to ritualized human culture. So we also

have a ‘down’ arrow from top to bottom of Table 1.

If social play and performance generate the necessary precondi-

tions for voluntary communication — increasing levels of self/other-

awareness and social trust — then we would expect an evolutionary

spiral with play and performance in one mode facilitating the emer-

gence of communication in a higher mode (Table 2). Even the simple

THE NEURAL CORRELATES OF WORK & PLAY 9

Communication Play Performance

Implicit Gesture-calls

(e.g. laughing,

crying)

Embodied

play (e.g.

contingent

mirror play)

Song-and-dance

display

Making marks

Mimetic Projective Iconic

gesture-calls

Projective

pretend play

Making

representational

images

Introjective Role-play Pantomime

Conventional Analogical codes

(e.g. pictographic

writing)

Collecting

behaviour

Ritual/ceremony

Iconography

Emblems

‘Fine art’

Music

Cryptic codes

(e.g. language,

phonetic

alphabets,

mathematical

denotations)

Play scripts Myth

Literary and

dramatic arts

Economico-moral

codes

Games-with-

rules

Socio-economic

personae

Wealth displays

(material,

moral, aesthetic,

cultural;

spiritual; etc.)

Table 1. Provisional classification showing some common forms of social

display (after Whitehead, 2001: based on data in Huizinga, 1955;

Bourdieu, 1972; Winnicott, 1974; Jennings, 1990, 1991; Burling, 1993).

mimetic signals of gorillas (Tanner and Byrne, 1996: in Mithen, 2005,

p. 118) imply a relatively high level of social insight by primate stan-

dards, presumably dependent on the playfulness of infant gorillas.

Song-and-dance display — by which I mean something comparable

with the synchronized balletic displays of dolphins (Connor, 1992) or

choral singing in gelada baboons (Richman, 1978, 1987) — fosters

shared experience in a potentially large number of participants, and

hence some ability to ‘put oneself in others’ shoes’. At the next higher

level, one would expect that ritualized mimetic performance would be

a precondition for the emergence of conventional communication —

notably language (Durkheim, 1912). Several arguments for this are

given in the Introduction to Knight (this volume).

Table 2 predicts three behavioural ‘rubicons’ during human evolu-

tion. The first, triggered by the emergence of song-and-dance display,

and the second, driven by major elaborations of pretend play, would

be expected to result in brain expansion. There are at least four rea-

sons why this might be so. Firstly, song-and-dance display involves

synchronized vocalization and movement with millisecond timing

precision (cf. Richman, 1978), and hence ‘massive neuronal

10 C. WHITEHEAD

Table 2. Hypothetical evolutionary sequence of social displays.

redundancy’ (Calvin, 1983).5 Secondly, all forms of play and perfor-

mance require multimodal sensorimotor integration (cf. Geschwind,

1967) and, thirdly, varying levels of skill (cf. Karni et al., 1995;

Turner and Whitehead, this volume). Fourthly, role-play requires the

ability to run multiple minds in parallel (see below).

The third rubicon — the emergence of ritualized culture — would

not be expected to create any selection pressure for brain expansion

and might even lead to some reduction in brain size. Brains are meta-

bolically and nutritionally expensive (Aiello and Wheeler, 1995), so

any reduction in selection pressure should lead to smaller brains.

Societies structured by rules and economico-moral systems may be

less dependent on precisely co-ordinated song-and-dance displays. If

this were the case, and if song-and-dance had been a factor in brain

expansion, then we would expect some reduction in brain size. It is

also likely that digital signals (such as language and scalar music)

would require less hardware than analogical ones (Brown, 1991; cf.

Knight, this volume), which might also contribute to some reduction

in brain size.

Does the fossil evidence support this? Figure 2 is a schematic repre-

sentation of hominid cranial capacities across a period of 3.5 million

years. This shows a pattern of punctuated equilibrium, with three

‘grade shifts’ or periods of accelerated change. In the first grade shift

cranial capacities doubled from an average of around 450 to 900 cm3.

In the second, average cranial capacity rose from 900 to 1500 cm3. So

far this is consistent with the hypothetical prediction. But what about

the third behavioural rubicon?

Human culture of modern type probably began in Africa before 100

kya (see Introduction to Knight, this volume). From the best available

data (De Miguel and Henneberg, 2001) it is difficult to say whether or

not brain expansion continued after the emergence of modern culture.

THE NEURAL CORRELATES OF WORK & PLAY 11

[5] Calvin (1983) showed that the ‘release window’ involved in throwing a missile at a targetcan be much shorter than the firing time of a single neurone (i.e. less than 1 msec). Thisrequires ‘massive neuronal redundancy’ to achieve the necessary precision. He calculatedthat hitting a 20 cm target at 14 m would require over 2,000 times more neurones than hit-ting the same target at 4 m. Earlier research by Richman (1978) had shown that geladababoons synchronize their voices with millisecond precision. He considered thisneuronally impossible unless individuals were predicting the vocalizations of others.However, in the case of human performance — for example that of a concert pianist — thesubtleties of rhythm, rubato, syncopation, and the characteristic ‘pulse’ that distinguishesthe work of different Western composers (Clines, 1977; Brown, 1991, p. 48) make multi-ple timing demands on both performer and audience. The enjoyment of music alsoinvolves ‘virtual dance’ affecting muscle tone throughout the body, especially the legs(Storr, 1993). Furthermore music both satisfies and deviates from audience expectations(Huron, 2006) so listeners cannot entirely predict the way the music is going — yet stillmaintain synchronized control over some very distributed neuronal systems.

However, there has certainly been a sharp decrease in cranial capacity

during the last ten thousand years — from an average of 1500 cm3

throughout the Upper Palaeolithic to 1350 cm3 in living humans. This

U-turn coincides with the agricultural revolution. One suggested

explanation is poor nutrition and the general decline in body size and

health that followed the switch from foraging to agriculture (Camilla

Power, personal communication). However, this does not explain why

body size recovered in well-fed modern nations, but brain size did not.

A second possible factor might be the rise of aristocratic systems and

centralized social control — without which no normal person would

have abandoned hunting for a grim life of toil, disease and malnutri-

tion. In contrast to egalitarian foragers, agricultural and industrial

populations are ‘controlled from the outside’ — and so presumably

are less dependent on integrative displays such as song-and-dance

performance. Research by Nicholas England (1968; 1995) revealed

that music does in fact correlate with political inequality. Egalitarian

and simple foraging peoples have free-flowing rhythm and melody,

whereas agrarian and more complex societies tend to have fixed

rhythmic and melodic structures. A more fluctuating performance

would presumably make more complex synchronization demands. I

12 C. WHITEHEAD

Figure 2. Schematic representation of three grade shifts affecting cranial

capacity during hominid evolution (based on data in Trinkaus and

Tompkins, 1990; Ruff et al., 1993; Aiello and Wheeler, 1995; Noble and

Davidson, 1996; De Miguel and Henneberg, 2001).

am not suggesting that living foragers — now displaced into the most

marginal environments — have larger brains than complex agri-

culturalists, but it is conceivable that different brain areas were rela-

tively more spared in different populations.

De Miguel and Henneberg (2001), whose data I have used in pre-

paring Figure 2, argue on statistical grounds that no ‘rubicons’ in

brain evolution can be inferred from cranial capacity data. Maciej

Henneberg, however, regards all hominids as a single lineage with no

more than one species living at any given time. Averaging robust apith

and habiline crania — despite anatomical evidence of diverging evo-

lutionary trajectories — obscures the first grade shift. Similarly,

lumping Homo erectus together with archaic Homo sapiens masks the

second grade shift. Further, cranial capacity data should not be viewed

in isolation. The apparent grade shifts coincide with structural changes

in brain anatomy and, in the early stages or just before each period of

change, there is archaeological evidence for major new forms of dis-

play (Whitehead, 2003; 2006b). I will give some examples below, but

first I want to consider recent brain imaging research which seems

consistent with a ‘play and display’ hypothesis of brain and self-

consciousness co-evolution.

Neurological Evidence for a ‘Play and Display’ Hypothesis

In Turner and Whitehead (this volume) we note that our brains contain

networks for representing the bodily actions of ourselves and others,

and areas that are involved with representing the ‘mental actions’ of

ourselves and others. We already have some evidence that both these

levels of mirroring ability exploit a similar principle (see Editorial,

this volume). In the studies which follow, we will see that hand actions

including the use of tools, unsurprisingly, activate the first level of

mirror networks. Dance also activates this level, plus some additional

areas. Projective pretend play (e.g. using a toy object to represent a

real object) activates a number of regions in common with dance,

along with the second level of mirroring areas — associated with

‘theory of mind’. Introjective pretend play (using yourself to repre-

sent another agent or object — i.e. role-play), I will argue on the basis

of a pilot study, activates all the pretend play areas and some addi-

tional ones which are also associated with creating and understanding

stories. We might think of these as ‘theatre of mind’ areas, and argu-

ably a distinct level of mirroring ability. All this suggests an evolu-

tionary sequence marked by a proliferation of ‘mirror systems’ in

successive hominid brains.

THE NEURAL CORRELATES OF WORK & PLAY 13

Imaging Studies of Pretend Play

I will begin ‘at the top’, with two kinds of pretend play. The first such

study, which we6 recently completed at the Wellcome Centre for

Imaging Neuroscience (Marchant et al., pending), exploited the fact

that familiar household objects can be used in three distinct ways —

the usual way (such as using a pen to write with), an unusual way

(such as using the pen to stir a cup of coffee), or a pretend way (such as

pretending that the pen is an aeroplane). We chose eighteen common

objects and prepared video sequences showing an actor using each

object in the three different ways.

A literature search revealed only one published study of pretence

(German et al., 2004). Previously, we7 had conducted a pilot study of

role-play (presented as a conference paper: Craik et al., 2000). So, to

my knowledge, there have only been three imaging studies of pretend

play.

Figure 3 (see colour section at end of this book) shows the major

centres of brain activity implicated by the three studies: that is, the

round markers are centred on the voxels (three-dimensional pixels)

showing the most significant peaks of activation. The square markers

represent a second task that formed part of our study — participants

were shown a picture of the object used in the preceding action video

and were required to name its use. Thus, in the case of the pen, they

might say ‘pen’, ‘coffee spoon’ or ‘aeroplane’. This was intended to

investigate brain areas involved in representing one object as another.

Notice that the earlier American study by German and colleagues

and our own study of pretence showed very similar patterns of activa-

tion. Both these studies involved people watching video clips of

actors pretending to do something, contrasted with videos of actors

doing non-pretend, instrumental actions — that is, brain activity asso-

ciated with observing non-pretend actions has been subtracted from

brain activity associated with observing pretence. The role-play study

shows a different pattern. Here, participants were not observing play,

but actually performing in imagination roles taken from Hamlet and

Macbeth. However, observing and performing generally involve com-

mon brain areas.

Figure 4 (see colour section at end of this book) shows a simplified

diagram of the main cortical areas activated by the two pretence

14 C. WHITEHEAD

[6] The research team comprised two neuroscientists — Jen Marchant and Chris Frith — andtwo social anthropologists — David Craik and Charles Whitehead.

[7] The role-play research team comprised Robert Turner (principal of the functional imaginglaboratory, Wellcome Centre for Imaging Neuroscience, at the time), David Craik andCharles Whitehead.

conditions — observing videos of pretence (projective) and mentally

performing role-play (introjective). This figure only shows the right

hemisphere of the brain, since all activations were either right-sided

or bilateral.

The first point to note here is that the temporal areas associated with

observing projective play (temperoparietal junction, superior tempo-

ral sulcus and temporal pole), and the medial prefrontal areas associ-

ated with both kinds of pretence, have been regularly associated with

‘theory of mind’ activity (Frith and Frith, 1999, 2000, 2001). ‘Theory

of mind’ (or ‘ToM’), otherwise known as mindreading or mentalizing

ability, is shorthand for the ability to interpret other people’s behav-

iour — and your own behaviour — in terms of mental states (Baron-

Cohen, 1995). Specifically, the term usually refers to epistemological

mental states — such as knowing, believing, pretending, imagining,

dreaming, etc. In other words, this is a particular level of self-

awareness and other-awareness, as distinct from the level of emotion

and desire.

German and colleagues, who did the earlier study of observing pre-

tence, took this to mean that ToM is automatically engaged whenever

people observe pretence. However, children engage in and recognize

pretend play two or three years before they show mindreading abili-

ties. This has led some authors to suggest that children have preco-

cious mentalizing abilities in the context of pretence (e.g. Leslie,

1987). There is considerable evidence against this — for example,

Angeline Lillard (1993) found that children can solve ToM tasks

before they appreciate that pretence involves mental as well as physi-

cal activity. Lillard’s (2001) ‘twin earth’ model concurs with social

mirror theory in making pretend play causally necessary for the devel-

opment of ToM, suggesting that the explanation is the other way

round — that is, ToM engages brain structures originally dedicated to

pretend play.

A second important point here is that all the areas indicated in

Figure 4 — whether associated with projective or introjective play —

coincide with areas regularly implicated in story-telling or narrative,

regardless of whether a story is told in words or in pictures (Mar,

2004). That is, the theory of mind network, together with the inferior

parietal lobule, dorsolateral prefrontal cortex, opercular prefrontal

cortex, and the posterior cingulate and precuneus, have all been impli-

cated in studies of narrative.

So ToM uses a subset of the brain areas implicated in two studies of

projective play, and projective play utilizes a subset of the areas

required for story-telling. We — that is, the research team involved in

THE NEURAL CORRELATES OF WORK & PLAY 15

the latest study of pretence [see note 6] — find it implausible that role-

play would not involve major ToM areas in the temporal lobes. The

fact that our earlier study of role-play did not show this, we think

implies that the control tasks in this study involved tacit role-play.8 If

that assumption is correct, then story-telling involves the same or very

similar brain areas to role-play. These results are open to more than

one possible interpretation, but if one accepts social mirror theory,

then a likely implication is that projective pretend play during child-

hood scaffolds both the development of role-play and ToM, and that

the ability to understand and enjoy stories involves pretend play

generally, including role-play.

Thirdly, all the cortices associated with projective play, role-play

and narrative — with the exception of the dorsolateral and opercular

frontal areas — are also commonly reported as so-called ‘deactivation

areas’. Cognitive neuroscientists have repeatedly observed that these

areas are more active during periods of supposed rest than they are

during laboratory tasks used to investigate cognitive processes

(Iacoboni et al., 2004). So researchers began to wonder what partici-

pants were doing when they were not supposed to be doing anything,

and some proposed that this resting brain activity was a ‘default state’

of the human brain. When we see that these areas coincide with activa-

tions associated with narrative, then it seems likely that people are

telling themselves stories during their ‘idle’ moments — in other

words, daydreaming.

Fourthly and finally I would like to point out something that

should not come as a surprise to anyone interested in the evolution of

the human brain. All the structures that we have discussed so far,

activated by make-believe, narrative, mindreading and probably day-

dreaming — in short, functions we might associate with imagina-

tion — are among the most expanded portions of human neocortex

(Whitehead, 2003).

The average human brain is around three times larger than the aver-

age chimp brain. However, not all human brain structures are equally

expanded relative to those of chimpanzees. Primary areas — such as

primary visual, primary motor and primary somatosensory areas —

are the same size in humans and chimps, because they do the same

kind of job. Secondary areas are two to three times larger in humans

relative to chimps. But the most massive expansions occurred in

16 C. WHITEHEAD

[8] All tasks were rehearsed and cued from rolling texts — Shakespearean script being con-trasted with legal and technical texts selected for their apparently uninvolving character.However, as Goffman (1959) would no doubt have agreed, professionalism means thatsuch texts are also likely to involve role-play.

multimodal integration areas. The prefrontal lobe is about six times

larger than a chimp’s, and the temporal lobe, especially the pole, is

also much larger. The inferior parietal lobule is also considerably

expanded — Norman Geschwind (1967) regarded this as ‘the impres-

sive advance’ in the human brain. This distinctive pattern of expan-

sions is consistent with social mirror theory and a ‘play and display’

hypothesis of human brain evolution.

One final comment on social imagination or ‘theatre of mind’

(Whitehead, 2001): to me, the truly amazing achievement of the

human brain is the ability to run social scenarios in imagination, with a

cast of actors — toy people — who behave as though they have minds,

knowledge, beliefs, desires and emotions of their own (Whitehead,

2001). The implication is that the human brain can run multiple

minds in parallel, and when this process goes wrong, the result is

dissociative identity disorder (cf. Hilgard et al., 1975; Bliss, 1986;

Brown, 1991; Mitchell, 1994).

Imaging Studies of Tool-Use and Object Manipulation

The industrial revolution and the protestant work ethic (Weber, 1904–5)

have created a world in which work is valued over play (Turner,

1982), object over social skills (Smith, 1988), logic over make-

believe (Jennings, 1990) and science and technology over the arts

(ibid). That may be one reason why we have more than forty imaging

studies of tool-use, object manipulation and hand action (Gr�zes and

Decety, 2001; footnote 9), as against only three studies of pretend

play.

All these manual action and manipulation studies followed from the

discovery of mirror neurones and were intended to investigate the

‘mirror system’ in human brains. Indeed the main areas activated are

considered to be homologues of mirror neurone areas in monkeys

(Gr�zes and Decety, 2001; Rizzolatti et al., 2001; Buccino et al.,

2001). Figure 5 (see colour section at end of this book) plots major

centres of brain activity from nine of these studies,9 seven of which

involved using and/or planning to use tools, whilst one involved nam-

ing tools (Martin et al., 1996) and the other hand-object interactive

movement (Naito et al., 2006). The first thing to note is that there are

major differences between tool/object manipulation and pretend play.

Firstly, brain activity associated with these instrumental tasks is more

THE NEURAL CORRELATES OF WORK & PLAY 17

[9] Martin et al. (1996); Choi et al. (2001); Inoui et al. (2001); Ohgami et al. (2004); Johnson-Frey et al. (2005); Fridman et al. (2006); Kan et al. (2006); Lotze et al. (2006); Naito et al.(2006).

in the left than the right hemisphere, whereas play involves more right

than left hemisphere activity. Secondly, there is a good deal of motor

and premotor involvement, as you would expect. Thirdly, there is a

cluster of superior parietal activations. This area of the brain is

involved with navigational body movements. Fourthly, there is very

little medial brain activity; and finally, very little prefrontal involve-

ment other than a large cluster of activations in the left frontal

operculum. Pretend play also involves this area, though mainly in the

right hemisphere.

The left prefrontal operculum includes Broca’s area, classically

linked to motor sequencing for speech. But it would seem that the area

is responsible for more than one kind of motor sequencing, including

grasping and using objects, playing with toys, role-play — and listen-

ing to stories even after the effects of language have been subtracted

out.

Imaging Studies of Dance

The left hemisphere pattern of activity associated with tool-use is

remarkably similar to that associated with dance, showing that dance,

too, involves the body-action mirror system, as one would expect

(Figure 6; see colour section at end of this book). Hence, dance

involves a great deal of activity in the opercular frontal area, a scatter

of motor activations, and a further cluster in the superior parietal lob-

ule. There is very little prefrontal activity other than in the operculum.

What is different about dance is that there is more right hemisphere

involvement, and also more medial activity — notably in the posterior

cingulate and precuneus — one area that we have associated with nar-

rative, role-play and daydreaming. Other areas common to pretend

play and narrative are the superior temporal sulcus and inferior pari-

etal lobule.

To the best of my knowledge there have been only four imaging

studies of dance, and Figure 6 shows the main activation loci from all

four. Further, three of these studies (Calvo-Merino et al., 2005, 2006;

Cross et al., 2006) were not intended to be studies of dance per se, but

were simply using dance as an alternative to object manipulation for

investigating mirror systems. Specifically, the authors wanted to

check whether knowing how to do something affected the way you

perceived others doing it. For example, the first of these studies

(Calvo-Merino et al., 2005) involved professional ballet dancers, pro-

fessional capoeira dancers, and non-dancer controls, watching videos

of ballet and capoeira. The results showed that when a dancer watches

18 C. WHITEHEAD

a dance form in which they have personal expertise mirror system

areas are more activated than when they watch a dance form that is not

familiar to them.

The lack of interest in the brain basis of dance is particularly odd

when you consider that entire research departments have been dedi-

cated to imaging studies of music, and there is a considerable volume

of neuroscientific literature devoted to performing and listening to

music. Further, studies of music are biased towards professional musi-

cians and classical music. Highly talented composers of classical

music in the West are regarded as ‘geniuses’, whereas highly talented

choreographers and non-classical composers are not, so this bias

seems to reflect the politics of prestige in Western culture. It may also

be easier to think of classical music as ‘cognitive’ relative to dance

and more ‘vulgar’ or embodied kinds of performance.

All the contrasts reported in both of the Calvo-Merino studies are,

in effect, dance minus dance — so they do not give a complete picture

of what is involved in observing dance. However, two of the four stud-

ies (Brown et al., 2006; Cross et al., 2006) did contrast dance tasks

against a resting baseline, although only the Cross et al. study gives a

visual map of this contrast (Figure 7; see colour section at end of this

book).

Evolution of the Brain and Self-Consciousness

This is all of potential palaeoanthropological interest, because the

most significant activations associated with dance are all in areas that

were expanded during the first phase of hominid brain expansion,

between 2.5 and 2 million years ago. The inferior parietal activations

reported in this study are of particular interest, though less marked

than one might have expected. A prominent inferior parietal lobule

first appears during this period of expansion (Tobias, 1987). When

you are dancing with other people there is an obvious need for

multimodal integration — you have to watch other dancers and listen

to any musical, vocal or percussive accompaniment, and integrate

your own movements to fit in with all these stimuli. That requires

visual, auditory, somatosensory and motor integration — which

implicates the inferior parietal lobule.

The finding that tool-use appears to engage a subset of brain areas

involved in dance is also of palaeoanthropological interest. The first

unequivocal knapped stone tools appear in the archaeological record

around 2.7 million years ago (Bilsborough, 1992, pp. 136–7), just

before the first period of major brain expansion. What is interesting

THE NEURAL CORRELATES OF WORK & PLAY 19

about these stone tools is not that they indicate any uniquely human

cognitive abilities (Wynn and McGrew, 1989). Orang-utans can learn

to make and use similar tools (Wright, 1972), and the bonobo chimp

Kanzi even invented new ways of making them (Toth et al., 1993).

What is immensely significant is that these tools were used for butch-

ering meat. When chimpanzees capture an animal they tear it apart

and eat it in a general m�lée (Teleki, 1973; 1981; Strum, 1981). They

are grabbing their share before the others eat it. Apes cannot afford the

luxury of butchering meat because they cannot trust each other to

share such a valuable resource. Butchering by these early hominids

indicates very high levels of social trust — unprecedented in non-

human apes. Further, the earliest butchery sites — at Hadar and Omo

in Ethiopia — are all beside rivers. If you are a prey animal, the last

thing you are likely to do is linger near fresh water, where dangerous

predators go to drink. Still less would you sit around knapping stone

tools and butchering meat, giving off an alluring smell that could

travel a long way downwind. The only hominids we know of at that

time were diminutive creatures little more than three feet tall. Yet they

apparently had no fear of dangerous social carnivores such as lions

and hyenas.

One possible explanation for these two facts — unprecedented

levels of social trust and the ability to drive off social carnivores much

larger than themselves — is song-and-dance display. Robin Dunbar

(1993) has shown that, in modern primates, there are straight-line

correlations between brain size, group size and time spent grooming

friends and allies. Projecting his findings onto fossil hominids

revealed that even the smallest-brained habiline — bearing in mind

how busy these early hominids were, carrying meat and stone to

butchery sites, making tools, butchering meat and confronting dan-

gerous carnivores — would not have had sufficient time for the

one-to-one grooming necessary to maintain the predicted group size

(Aiello and Dunbar, 1993). Dunbar postulated ‘vocal grooming’ —

allowing multiple individuals to be groomed simultaneously — as a

likely origin for language. Anthropological, palaeogenetic and chrono-

linguistic arguments would rule out language at this early date (see

Introduction to Knight, this volume), but ‘song’ would in any case

equate better with ‘vocal grooming’. I have discussed this in more

detail elsewhere along with other evidence (Whitehead, 2006b). Suf-

fice it to say that the above imaging research is consistent with the

possibility that song-and-dance display was a significant factor driv-

ing the first phase of hominid brain expansion.

20 C. WHITEHEAD

It also seems likely that song-and-dance display would create the

necessary preconditions for the later emergence of sophisticated

mimetic displays — increased social trust, social insight and volun-

tary control over display behaviours (Whiten, 1993). Dance may also

have pre-adapted the brain for mimetic displays such as iconic signals,

mime, pretend play and role-play. We would expect song-and-dance

to lead to expansions of inferior parietal and opercular prefrontal

areas, together with temperoparietal and superior temporal areas and

the posterior cingulate gyrus and precuneus, which are also implicated

in pretend play and role-play. Whilst fossil crania show no clear evi-

dence of prefrontal expansion across the first phase of accelerated

changes in the brain — other than the opercular prefrontal area impli-

cated in dance and hand actions — some expansion of ventromedial

prefrontal cortex is a possibility. This could be regarded as a ‘groom-

ing centre’ — it is involved in feelings of empathy and response to

pleasurable bodily contact, pleasant words and harmonious as opposed

to discordant music. The functions of dance of course include groom-

ing and entrainment (Whitehead, 2001; 2003).

However, the major expansion of the prefrontal lobe — which

gives us humans our vertical forehead and ‘highbrow’ appearance —

did not occur until the second phase of hominid brain expansion. This

coincided with the first archaeological evidence for new forms of dis-

play — the use of red pigment, collections of attractive objects, and

possibly the world’s oldest doll, the Berekhat Ram figurine, which is

over 230,000 years old. The increasing ‘playfulness’ of the archaeo-

logical record during this period might suggest that frontal lobe

expansion was associated with increasingly sophisticated pretend

play, although this period also saw the descent of the human larynx to

create a large tuneable pharynx, suggesting melodic elaboration of

song (Whitehead, 2006b).

Conclusion

All the evidence I have presented is consistent with social mirror

theory and a ‘play and display’ hypothesis of human brain expansion.

According to social mirror theory, different modes of display create

different levels of self-consciousness and social insight. Implicit dis-

plays such as affective gesture-calls, embodied play and song-and-

dance lead to insight at the level of emotion, desire and relationships.

They are the basis of empathy. Mimetic displays such as pretend play

and role-play lead to insight at the level of epistemological mental

states — that is, theory of mind and theatre of mind. Such displays are

THE NEURAL CORRELATES OF WORK & PLAY 21

part of our biological heritage, and they are the foundations on which

conventional displays are built — ritual, language, music and wealth

displays — which characterize and constitute modern human culture.

However, our ancestors achieved modern-type culture at a price.

Conventional displays do not simply create a new mode of self- and

other-consciousness with an increased capacity for abstract thought.

They also produce a kind of unconsciousness. It is the job of culture to

falsify our self/other perceptions and conceal our biologically-given

nature. This was the only way that our social but selfish ancestors

could be coerced into collaborating in a radically unselfish and anti-

biological system.

The result is collective deception. Collective deceptions in Western

science include physicalism, individualism, cognocentrism, logo-

centrism, genocentrism — and the cultural materialism of the West,

which regards work as the opposite of play, and play as basically a

waste of time. The scientific method of course is designed to obviate

such biases and scientists are already beginning to recognize them.

Some of the authors in this volume have been studying social displays

for decades, which would put them in the vanguard of a trend which,

for reasons given above, I think will expand the horizons of research

into human behaviour.

Whilst conducting a literature search on pretend play, I was struck

by the recurring admissions of difficulty in defining play. Authors reg-

ularly resort at some point to the phrase ‘just for fun’ — in quotes

because they are resorting to folk terminology. There is no scientific

definition of ‘fun’ — yet it is surely self-evident that when we are hav-

ing fun, that is when we are being most true to our biologically given

nature, and when the brain and the body are doing their most natural

work. It is high time we saw a lot more research on the displays, art-

istry and creative imagination that make us most truly human.

Acknowledgements

I am grateful to Professors Robert Turner and Chris Frith for gener-

ously sharing their research funding, making available the resources

of the Wellcome Trust Centre for Neuroimaging, and for encouraging,

supporting and supervising the studies of role-play and pretend play

reported here. I would also like to thank Dr Jen Marchant for doing

most of the work and analysis on the pretend play study, and others at

the Wellcome Trust Centre for help and advice on countless occa-

sions. I am also grateful to David Craik whose long experience in

theatre, cinema and television was vital to both studies, Professor Dr

22 C. WHITEHEAD

Maciej Henneberg for providing his cranial capacity data, Professor

Emily Cross for providing the image shown as Figure 7, and my son

Jon James for redrawing the cartoon sequence which Warner Brothers

could not identify. Imaging research was funded by the Wellcome

Trust.

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