Brains can tell us more about social cognition if our methods don’t presuppose the answers.

Ian Apperly

Brains can tell us more about social cognition the cognitive basis of “theory of mind”if our methods don’t presuppose the answers.

Ian Apperly

What is “Theory of Mind”?• “Folk psychology”, “Perspective-taking”, “Social cognition”• Essential for everyday social interaction and communication

• False belief tasks as a paradigm case• (e.g., Wimmer & Perner, 1983)

– These tasks ensure that participant must judge from other person’s point of view

What is “Theory of Mind”?• “Folk psychology”, “Perspective-taking”, “Social cognition”• Essential for everyday social interaction and communication

• False belief tasks as a paradigm case• (e.g., Wimmer & Perner, 1983)

– These tasks ensure that participant must judge from other person’s point of view

• Significant developments from infancy to early childhood

• Disproportionately impaired in autism and several other genetic and psychiatric disorders

What is “Theory of Mind”?• “Folk psychology”, “Perspective-taking”, “Social cognition”• Essential for everyday social interaction and communication

• False belief tasks as a paradigm case• (e.g., Wimmer & Perner, 1983)

– These tasks ensure that participant must judge from other person’s point of view

• Significant developments from infancy to early childhood

• Disproportionately impaired in autism and several other genetic and psychiatric disorders

• Existent, to a degree, in non-human animals

What is “Theory of Mind”?• “Folk psychology”, “Perspective-taking”, “Social cognition”• Essential for everyday social interaction and communication

• False belief tasks as a paradigm case• (e.g., Wimmer & Perner, 1983)

– These tasks ensure that participant must judge from other person’s point of view

• Significant developments from infancy to early childhood

• Disproportionately impaired in autism and several other genetic and psychiatric disorders

• Existent, to a degree, in non-human animals

• Identifiable neural networkTemporo-parietal junction / pSTS

Temporal poleMedial prefrontal cortex

Lateral view

TPJ

TP

Medial view

mPF

C

Theory of mind in adults?

• “But don’t adults have a theory of mind……?”

Theory of mind in adults?

• “But don’t adults have a theory of mind……?”

• Prevailing view:– ToM is a set of concepts– Researchers should figure out who has them (and where

they are in the brain).....– ....by seeing who passes false belief tasks

Temporo-parietal junction / pSTSTemporal poleMedial prefrontal cortex

Lateral view

TPJ

TP

Medial view

mPF

C

Theory of mind in adults?

• “But don’t adults have a theory of mind……?”

• Prevailing view:– ToM is a set of concepts– Researchers should figure out who has them (and where

they are in the brain).....– ....by seeing who passes false belief tasks

• Problems with this view:– No cognitive account of ToM in adults– Severe limitations on conceptualising extended

development, neural basis and disorder – Little integration with the rest of cognition

Temporo-parietal junction / pSTSTemporal poleMedial prefrontal cortex

Lateral view

TPJ

TP

Medial view

mPF

C

Background: The “theory of mind network”

Temporo-parietal junction

Temporal pole

Medial prefrontal cortex

Left lateral view

TPJ

TP

Medial view

mPF

C

Anterior Posterior

PC TPJ

TPmPFC

Precuneus

TPJ

TP

Anterior

PC

Right lateral view

e.g. Frith & Frith, 2003 Van Overwalle, 2009

Background: The “theory of mind network”

Temporo-parietal junction

Temporal pole

Medial prefrontal cortex

Left lateral view

TPJ

TP

Medial view

mPF

C

Anterior Posterior

PC TPJ

TPmPFC

Precuneus

TPJ

TP

Anterior

PC

Right lateral view

e.g. Frith & Frith, 2003 Van Overwalle, 2009

Background: The “theory of mind network”

Temporo-parietal junction

Temporal pole

Medial prefrontal cortex

Left lateral view

TPJ

TP

Medial view

mPF

C

Anterior Posterior

PC TPJ

TPmPFC

Precuneus

TPJ

TP

Anterior

PC

Right lateral view

e.g. Frith & Frith, 2003 Van Overwalle, 2009

Main debate is around which regions are “really” ToM regions – i.e. Where is the ToM module?

ToM functional localiser(Saxe & Kanwisher, 2003......)

False belief (FB) sample storyJohn told Emily that he had a Porsche.Actually, his car is a Ford. Emilydoesn’t know anything about carsthough, so she believed John.—When Emily sees John’s car shethinks it is aporsche ford

False photograph (FP) sample storyA photograph was taken of an apple hangingon a tree branch. The film took half an hour todevelop. In the meantime, a strongwind blew the apple to the ground.—The developed photograph shows the apple on theground branch

ToM functional localiser(Saxe & Kanwisher, 2003......)

False belief (FB) sample storyJohn told Emily that he had a Porsche.Actually, his car is a Ford. Emilydoesn’t know anything about carsthough, so she believed John.—When Emily sees John’s car shethinks it is aporsche ford

False photograph (FP) sample storyA photograph was taken of an apple hangingon a tree branch. The film took half an hour todevelop. In the meantime, a strongwind blew the apple to the ground.—The developed photograph shows the apple on theground branch

R-TPJ shows greatest specificity for reasoning about mental states. Contrast with mPFC, which also shows activity for thinking about body states, internal sensations and personal characteristics.

So is this the ToM module?

Why ToM cannot be a Fodor-module

Why ToM cannot be a Fodor-module

• According to Fodor (1983, 2000) deciding what we believe is an archetypal “central” process

?

Why ToM cannot be a Fodor-module

• According to Fodor (1983, 2000) deciding what we believe is an archetypal “central” process

• It would be odd, in the extreme, if deciding what we believed someone else believed were somehow modular

??

What might we expect Mindreading to involve?

What might we expect Mindreading to involve?

Well, I was caned in my time and I’ve concentrated all my life

Do you not think, Sir Rhodes, if you get caned in school

you can’t concentrate?

You was caned? Respect man,

respect

What might we expect Mindreading to involve?

• Conceptual knowledge about mental states

• Represent alternative perspectives

Well, I was caned in my time and I’ve concentrated all my life

Do you not think, Sir Rhodes, if you get caned in school

you can’t concentrate?

You was caned? Respect man,

respect

What might we expect Mindreading to involve?

• Conceptual knowledge about mental states

• Represent alternative perspectives• Keep up!• Avoid interference from self

perspective

Well, I was caned in my time and I’ve concentrated all my life

Do you not think, Sir Rhodes, if you get caned in school

you can’t concentrate?

You was caned? Respect man,

respect

What might we expect Mindreading to involve?

• Conceptual knowledge about mental states

• Represent alternative perspectives• Keep up!• Avoid interference from self

perspective• Make abductive, “best guess”

inferences • Do this in the context of relevant

social scriptsWell, I was caned in my time and I’ve concentrated all my life

Do you not think, Sir Rhodes, if you get caned in school

you can’t concentrate?

You was caned? Respect man,

respect

Neuroimaging studies that are starting to cast light on these functions, and their neural correlates

Belief-desire reasoning

• Young children pass true belief tasks (~3Y) before false belief tasks (~4Y) (e.g., Bartsch & Wellman, 1988)

Diffi

culty

B+ B-

True belief

False belief

Belief-desire reasoning

• Young children pass true belief tasks before false belief tasks (e.g., Bartsch & Wellman, 1988)

• Young children pass false belief tasks at ~4 years when protagonist wishes to find object, but not until ~5 years when protagonist wishes to avoid object (e.g., Cassidy, 1998; Friedman & Leslie, 2004)

True belief

False belief

Diffi

culty

B+ B-

D-

D+

Orthogonal variation of beliefs and desires(Hartwright, Apperly & Hansen, 2012)

Behavioural study(Apperly et al., 2011, Ch.Dev.;

0

0.1

0.2

0.3

0.4

0.5

0.6

B+ B- B+ B- B+ B- B+ B-

6-7Y 8-9Y 10-11Y Adults Adults

Belief

Pro

po

rtio

n o

f err

ors

D+

D-

0

500

1000

1500

2000

2500

3000

3500

B+ B- B+ B- B+ B- B+ B-

6-7Y 8-9Y 10-11Y Adults Adults

Belief

Res

pons

e Ti

me

(ms)

D+

D-

Children’s dataApperly, Warren, et al. (2012)

RT to correct responses Errors

Main Effects: Belief, Desire, AgeAge*Desire – but Desire significant at all ages

Diffi

culty

B+ B-

D-D+

0

0.1

0.2

0.3

0.4

0.5

0.6

B+ B- B+ B- B+ B- B+ B-

6-7Y 8-9Y 10-11Y Adults Adults

Belief

Pro

po

rtio

n o

f err

ors

D+

D-

0

500

1000

1500

2000

2500

3000

3500

B+ B- B+ B- B+ B- B+ B-

6-7Y 8-9Y 10-11Y Adults Adults

Belief

Res

pons

e Ti

me

(ms)

D+

D-

Children’s dataApperly, Warren, et al. (2012)

RT to correct responses Errors

Main Effects: Belief, Desire, AgeAge*Desire – but Desire significant at all ages

Main Effects: Belief, Desire, AgeAge*Desire –Desire significant only at 6-7 and 8-9

Diffi

culty

B+ B-

D-D+

0

0.1

0.2

0.3

0.4

0.5

0.6

B+ B- B+ B- B+ B- B+ B-

6-7Y 8-9Y 10-11Y Adults Adults

Belief

Pro

po

rtio

n o

f err

ors

D+

D-

0

500

1000

1500

2000

2500

3000

3500

B+ B- B+ B- B+ B- B+ B-

6-7Y 8-9Y 10-11Y Adults Adults

Belief

Res

pons

e Ti

me

(ms)

D+

D-

Adults’ data

RT to correct responses Errors

Diffi

culty

B+ B-

D-D+

Belief, DesireBelief*Desire – all comparisons significant

Consistent with German & Hehman (2006)

0

0.1

0.2

0.3

0.4

0.5

0.6

B+ B- B+ B- B+ B- B+ B-

6-7Y 8-9Y 10-11Y Adults Adults

Belief

Pro

po

rtio

n o

f err

ors

D+

D-

0

500

1000

1500

2000

2500

3000

3500

B+ B- B+ B- B+ B- B+ B-

6-7Y 8-9Y 10-11Y Adults Adults

Belief

Res

pons

e Ti

me

(ms)

D+

D-

Adults’ data

RT to correct responses Errors

Diffi

culty

B+ B-

D-D+

Belief, DesireBelief*Desire – all comparisons significant

Belief, not Desire

Consistent with German & Hehman (2006)

Orthogonal variation of beliefs and desires(Hartwright, Apperly & Hansen, 2012)

Behavioural study(Apperly et al., 2011, Ch.Dev.;

Orthogonal variation of beliefs and desires(Hartwright, Apperly & Hansen, 2012)

• B- is harder than B+

• D- is harder than D+

• This replicates findings from children and adults

– (Apperly et al., 2011, Ch.Dev.;

Belief (True vs. False) TPJ, ACC, IFGDesire (Like vs. Hate) TPJ, ACC

Overlap

Orthogonal variation of beliefs and desires(Hartwright, Apperly & Hansen, 2012)

Belief (True vs. False) TPJ, ACC, IFGDesire (Like vs. Hate) TPJ, ACC

Overlap

Orthogonal variation of beliefs and desires(Hartwright, Apperly & Hansen, 2012)

Notably no mPFC

Belief-desire task vs. ToM-localiser

Belief OR Desire“ToM localiser” (False Belief – False Photo)OverlapConjunction analysis between Belief-Desire and ToM Localiser

Orthogonal variation of beliefs and desires(Hartwright, Apperly & Hansen, 2012)

• Varying Belief and Desire (not belief or desire per se) modulates activity in– “control” areas (ACC) – perhaps reflecting variation in condition difficulty– “ToM” areas (bilateral TPJ)

Orthogonal variation of beliefs and desires(Hartwright, Apperly & Hansen, 2012)

• Varying Belief and Desire (not belief or desire per se) modulates activity in– “control” areas (ACC) – perhaps reflecting variation in condition difficulty– “ToM” areas (bilateral TPJ)

• Varying Belief (but not Desire) modulates– “control” areas (IFG – R-IFG in particular) – only B- vs. B+ involves a perspective

difference

Orthogonal variation of beliefs and desires(Hartwright, Apperly & Hansen, 2012)

• Varying Belief and Desire (not belief or desire per se) modulates activity in– “control” areas (ACC) – perhaps reflecting variation in condition difficulty– “ToM” areas (bilateral TPJ)

• Varying Belief (but not Desire) modulates– “control” areas (IFG – R-IFG in particular) – only B- vs. B+ involves a perspective

difference

• Why are “control” areas not observed in ToM localiser?– False Photo subtracts this from False Belief

Orthogonal variation of beliefs and desires(Hartwright, Apperly & Hansen, 2012)

• Varying Belief and Desire (not belief or desire per se) modulates activity in– “control” areas (ACC) – perhaps reflecting variation in condition difficulty– “ToM” areas (bilateral TPJ)

• Varying Belief (but not Desire) modulates– “control” areas (IFG – R-IFG in particular) – only B- vs. B+ involves a perspective

difference

• Why are “control” areas not observed in ToM localiser?– False Photo subtracts this from False Belief

• Why is mPFC observed in localiser but not our task?– Our task does not require abductive “uncertain” inferences

Social abduction(Hartwright, Apperly & Hansen, in prep)

Social abduction(Hartwright, Apperly & Hansen, subm.)

Selective for D?

TB vs. FB

Green = D? vs. D-&D+

Green = D? vs. D-&D+&FB&TB

Case study 2 – Temporal coordination

Background: The neural basis of “theory of mind”

Temporo-parietal junction

Temporal pole

Medial prefrontal cortex

Left lateral view

TPJ

TP

Medial view

mPF

C

Anterior Posterior

PC TPJ

TPmPFClPFC

Precuneus

TPJ

TP

Anterior

lPFClPFC

PCLateral prefrontal cortex

Right lateral view

We don’t know how these regions work together

e.g. Frith & Frith, 2003 Van Overwalle, 2009

Automatic perspective-taking?(Samson, Apperly, Braithwaite et al., 2010, JEP:HPP)

You / He

You / He

2

2

Self / Other Consistent

Self / Other InconsistentDisc position varies

1,2, or 3 discs

You / He

You / He

2

2

Self / Other Consistent

Self / Other InconsistentDisc position varies

1,2, or 3 discs

Automatic perspective-taking?(Samson, Apperly, Braithwaite et al., 2010, JEP:HPP)

Main effect of consistencySignificant interaction

400

450

500

550

600

650

700

750

800

850

900

Self Other Self Other Self Other

Discs vary Figure varies Blocked

Consistent

InconsistentRT (m

s)

Egocentric interference on explicit judgement of other

Automatic perspective-taking?(Samson, Apperly, Braithwaite et al., 2010, JEP:HPP)

Main effect of consistencySignificant interaction

400

450

500

550

600

650

700

750

800

850

900

Self Other Self Other Self Other

Discs vary Figure varies Blocked

Consistent

InconsistentRT (m

s)

Altercentric interference =evidence of automatic calculation of perspective

Automatic perspective-taking?(Samson, Apperly, Braithwaite et al., 2010, JEP:HPP)

Main effect of consistencySignificant interaction

400

450

500

550

600

650

700

750

800

850

900

Self Other Self Other Self Other

Discs vary Figure varies Blocked

Consistent

InconsistentRT (m

s)

Various follow-ups.....

Altercentric interference = evidence of automatic calculation of perspective

Automatic perspective-taking?(Samson, Apperly, Braithwaite et al., 2010, JEP:HPP)

Automatic perspective-taking?(Samson, Apperly, Braithwaite et al., 2010, JEP:HPP)

Only ever judge “self” – how many dots you can see

500

550

600

650

700

750

800

Self - avatar distractor Self - rectangle distractor

Experiment 3R

eact

ion

tim

e (m

s)

Consistent

Inconsistent* ns

Automatic perspective-taking?(Samson, Apperly, Braithwaite et al., 2010, JEP:HPP)

Only ever judge “self” – how many dots you can see

Automatic and controlled processes within a perspective-taking problem?

Main effect of consistencySignificant interaction

400

450

500

550

600

650

700

750

800

850

900

Self Other Self Other Self Other

Discs vary Figure varies Blocked

Consistent

InconsistentRT (m

s)

Altercentric interference = indication of automatic perspective calculation

Calculation Selection Response

SelfSelf Yes

Other

Automatic and controlled processes within a perspective-taking problem?

Main effect of consistencySignificant interaction

400

450

500

550

600

650

700

750

800

850

900

Self Other Self Other Self Other

Discs vary Figure varies Blocked

Consistent

InconsistentRT (m

s)

Altercentric interference = indication of automatic perspective calculation

Dual tasking

Calculation Selection Response

SelfSelf Yes

Other

Cognitively effortful perspective selectionQureshi, Apperly & Samson (2010) Cognition.

0

200

400

600

800

1000

1200

Alone Dual Alone Dual

Other Self

Pro

cess

ing

co

sts

(RT

/pro

p.

corr

ect)

Consistent

Inconsistent

Altercentric interference is increased by dual tasking with an executive task

Background: The neural basis of “theory of mind”

Temporo-parietal junction

Temporal pole

Medial prefrontal cortex

Left lateral view

TPJ

TP

Medial view

mPF

C

Anterior Posterior

PC TPJ

TPmPFClPFC

Precuneus

TPJ

TP

Anterior

lPFClPFC

PCLateral prefrontal cortex

Right lateral view

e.g. Frith & Frith, 2003 Van Overwalle, 2009

Background: The neural basis of “theory of mind”

Temporo-parietal junction

Temporal pole

Medial prefrontal cortex

Left lateral view

TPJ

TP

Medial view

mPF

C

Anterior Posterior

PC TPJ

TPmPFClPFC

Precuneus

TPJ

TP

Anterior

lPFClPFC

PCLateral prefrontal cortex

Right lateral view

e.g. Frith & Frith, 2003 Van Overwalle, 2009

Background: The neural basis of “theory of mind”

Temporo-parietal junction

Temporal pole

Medial prefrontal cortex

Left lateral view

TPJ

TP

Medial view

mPF

C

Anterior Posterior

PC TPJ

TPmPFClPFC

Precuneus

TPJ

TP

Anterior

lPFClPFC

PCLateral prefrontal cortex

Right lateral view

e.g. Frith & Frith, 2003 Van Overwalle, 2009

Background: The neural basis of “theory of mind”

Temporo-parietal junction

Temporal pole

Medial prefrontal cortex

Left lateral view

TPJ

TP

Medial view

mPF

C

Anterior Posterior

PC TPJ

TPmPFClPFC

Precuneus

TPJ

TP

Anterior

lPFClPFC

PCLateral prefrontal cortex

Right lateral view

We don’t know how these regions work together

e.g. Frith & Frith, 2003 Van Overwalle, 2009

Predictions for an ERP study

• Functionally, we have evidence for an initial process of perspective calculation followed by a later process of perspective selection

• Calculation: Where do we first see discrimination between Self and Other conditions? (Anterior/Frontal versus Posterior/Temporo-parietal)

• Selection: Predict later process in lPFC (perhaps right lPFC), that differentiates Congruent and Incongruent conditions.

ERP study(McCleery et al., 2011, Journal of Neuroscience)

• Pilot study (N=8) identified electrode sets in which we observed differentiation of conditions.

• Main study (N=17) 192 trials per condition• Behavioural effects

– Self<Other in RTs– Consistent<Inconsistent in RTs and Errors– Effect of Consistency was greatest for Other

• ERP recorded from onset of picture

Perspective calculation:450ms Self<Other latency over posterior scalp

Confirmatory source analysis suggested Bilateral TPJ

Perspective selection:LSW (600-800ms) Inconsistent<Consistent amplitude over right anterior scalp

Perspective selection:LSW (600-800ms) Inconsistent<Consistent amplitude over right anterior scalp

Right inferior frontal gyrus was the only source to discriminate Inconsistent<Consistent for both Self and Other

ConclusionsPrimacy for posterior regions in perspective calculation – at least for simple

perspectives

Temporo-parietal junction

Temporal pole

Medial prefrontal cortex

Left lateral view

TPJ

TP

Medial view

mPF

C

Anterior Posterior

PC TPJ

TPmPFClPFC

Precuneus

TPJ

TP

Anterior

lPFClPFC

PCLateral prefrontal cortex

Right lateral view

e.g. Frith & Frith, 2003 Van Overwalle, 2009

ConclusionsRole for non-ToM “control network” in perspective selection

Temporo-parietal junction

Temporal pole

Medial prefrontal cortex

Left lateral view

TPJ

TP

Medial view

mPF

C

Anterior Posterior

PC TPJ

TPmPFClPFC

Precuneus

TPJ

TP

Anterior

lPFClPFC

PCLateral prefrontal cortex

Right lateral view

e.g. Frith & Frith, 2003 Van Overwalle, 2009

What might we expect Mindreading to involve?The “ToM network”

• Conceptual knowledge about mental states

• Represent alternative perspectives• Keep up!• Avoid interference from self

perspective• Make abductive, “best guess”

inferences • Do this in the context of relevant

social scripts

Well, I was caned in my time and I’ve concentrated all my life

Do you not think, Sir Rhodes, if you get caned in

school you can’t concentrate?

You was caned? Respect man,

respect

TPJ

TP

Right lateral view

Medial view

mPF

C

PC

• Conceptual knowledge about mental states

• Represent alternative perspectives• Keep up!• Avoid interference from self

perspective• Make abductive, “best guess”

inferences • Do this in the context of relevant

social scripts

Well, I was caned in my time and I’ve concentrated all my life

Do you not think, Sir Rhodes, if you get caned in

school you can’t concentrate?

You was caned? Respect man,

respect

TPJ

TP

Right lateral view

Medial view

mPF

C

PC

What might we expect Mindreading to involve?The “ToM network”

• Conceptual knowledge about mental states

• Represent alternative perspectives• Keep up!• Avoid interference from self

perspective• Make abductive, “best guess”

inferences • Do this in the context of relevant

social scripts ?????????

Well, I was caned in my time and I’ve concentrated all my life

Do you not think, Sir Rhodes, if you get caned in

school you can’t concentrate?

You was caned? Respect man,

respect

TPJ

TP

Right lateral view

Medial view

mPF

C

PC

What might we expect Mindreading to involve?The “ToM network”

• Conceptual knowledge about mental states

• Represent alternative perspectives• Keep up!• Avoid interference from self

perspective• Make abductive, “best guess”

inferences • Do this in the context of relevant

social scripts

Well, I was caned in my time and I’ve concentrated all my life

Do you not think, Sir Rhodes, if you get caned in

school you can’t concentrate?

You was caned? Respect man,

respect

TPJ

TP

Right lateral view

Medial view

mPF

C

PCACC

What might we expect Mindreading to involve?Cognitive control

• Conceptual knowledge about mental states

• Represent alternative perspectives• Keep up!• Avoid interference from self

perspective• Make abductive, “best guess”

inferences • Do this in the context of relevant

social scripts

Well, I was caned in my time and I’ve concentrated all my life

Do you not think, Sir Rhodes, if you get caned in

school you can’t concentrate?

You was caned? Respect man,

respect

TPJ

TP

lPFC

Right lateral view

Medial view

mPF

C

PC

What might we expect Mindreading to involve?Cognitive control

ACC

• Conceptual knowledge about mental states

• Represent alternative perspectives• Keep up!• Avoid interference from self

perspective• Make abductive, “best guess”

inferences • Do this in the context of relevant

social scripts

• ???Well, I was caned in my time and I’ve concentrated all my life

Do you not think, Sir Rhodes, if you get caned in

school you can’t concentrate?

You was caned? Respect man,

respect

TPJ

TP

lPFC

Right lateral view

Medial view

mPF

C

PC

What might we expect Mindreading to involve?Cognitive control

ACC

• Conceptual knowledge about mental states• Represent alternative perspectives• Keep up!• Avoid interference from self perspective• Make abductive, “best guess” inferences • Do this in the context of relevant social

scripts

• Whether or not these particulars are correct.....

• “Where is the ToM module” is a poorly conceived question

• Functional and neural studies are combining to give new insights into what ToM is, and how we do it.

Well, I was caned in my time and I’ve concentrated all my life

Do you not think, Sir Rhodes, if you get caned in

school you can’t concentrate?

You was caned? Respect man,

respect

TPJ

TP

lPFC

Right lateral view

Medial view

mPF

C

PC

What might we expect Mindreading to involve?

ACC

Orthogonal variation of mental/non-mental and ambiguous/unambiguous inferences(Jenkins & Mitchell, 2009, Cereb.Cortex.)

Orthogonal variation of mental/non-mental and ambiguous/unambiguous inferences(Jenkins & Mitchell, 2009, Cereb.Cortex.)

Main effect of Mental/non-mental in rTPJMain effect of ambiguous/unambiguous in mPFC

Social abduction(Hartwright, Apperly & Hansen, in prep)