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

2. 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

3. 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

4. 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

5. 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

6. 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 network Temporo-parietal junction / pSTS Temporal pole Medial prefrontal cortex Lateral view TPJ TP Medial view mPFC

7. Theory of mind in adults? “But don’t adults have a theory of mind……?”

8. 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 / pSTS Temporal pole Medial prefrontal cortex Lateral view TPJ TP Medial view mPFC

9. 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 / pSTS Temporal pole Medial prefrontal cortex Lateral view TPJ TP Medial view mPFC

10. Background: The “theory of mind network” Temporo-parietal junction Temporal pole Medial prefrontal cortex Left lateral view TPJ TP Medial view mPFC AnteriorPosterior PC TPJ TP mPFC Precuneus TPJ TP Anterior PC Right lateral view e.g. Frith & Frith, 2003 Van Overwalle, 2009

11. Background: The “theory of mind network” Temporo-parietal junction Temporal pole Medial prefrontal cortex Left lateral view TPJ TP Medial view mPFC AnteriorPosterior PC TPJ TP mPFC Precuneus TPJ TP Anterior PC Right lateral view e.g. Frith & Frith, 2003 Van Overwalle, 2009

12. Background: The “theory of mind network” Temporo-parietal junction Temporal pole Medial prefrontal cortex Left lateral view TPJ TP Medial view mPFC AnteriorPosterior PC TPJ TP mPFC 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?

13. ToM functional localiser (Saxe & Kanwisher, 2003......) False belief (FB) sample story John told Emily that he had a Porsche. Actually, his car is a Ford. Emily doesn’t know anything about cars though, so she believed John. — When Emily sees John’s car she thinks it is a porsche ford False photograph (FP) sample story A photograph was taken of an apple hanging on a tree branch. The film took half an hour to develop. In the meantime, a strong wind blew the apple to the ground. — The developed photograph shows the apple on the ground branch

14. ToM functional localiser (Saxe & Kanwisher, 2003......) False belief (FB) sample story John told Emily that he had a Porsche. Actually, his car is a Ford. Emily doesn’t know anything about cars though, so she believed John. — When Emily sees John’s car she thinks it is a porsche ford False photograph (FP) sample story A photograph was taken of an apple hanging on a tree branch. The film took half an hour to develop. In the meantime, a strong wind blew the apple to the ground. — The developed photograph shows the apple on the ground 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?

15. Why ToM cannot be a Fodor-module

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

17. 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 ? ?

18. What might we expect Mindreading to involve?

19. 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

20. 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

21. 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

22. 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 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

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

24. Belief-desire reasoning Young children pass true belief tasks (~3Y) before false belief tasks (~4Y) (e.g., Bartsch & Wellman, 1988) Difficulty B+ B- True belief False belief

25. 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 Difficulty B+B- D- D+

26. Orthogonal variation of beliefs and desires (Hartwright, Apperly & Hansen, 2012) Behavioural study (Apperly et al., 2011, Ch.Dev.;

27. Children’s data Apperly, Warren, et al. (2012) RT to correct responses Errors Main Effects: Belief, Desire, Age Age*Desire – but Desire significant at all ages Difficulty B+B- D- D+

28. Children’s data Apperly, Warren, et al. (2012) RT to correct responses Errors Main Effects: Belief, Desire, Age Age*Desire – but Desire significant at all ages Main Effects: Belief, Desire, Age Age*Desire –Desire significant only at 6-7 and 8-9 Difficulty B+B- D- D+

29. Adults’ data RT to correct responses Errors Difficulty B+B- D- D+ Belief, Desire Belief*Desire – all comparisons significant Consistent with German & Hehman (2006)

30. Adults’ data RT to correct responses Errors Difficulty B+B- D- D+ Belief, Desire Belief*Desire – all comparisons significant Belief, not Desire Consistent with German & Hehman (2006)

31. Orthogonal variation of beliefs and desires (Hartwright, Apperly & Hansen, 2012) Behavioural study (Apperly et al., 2011, Ch.Dev.;

32. 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.;

33. Belief (True vs. False) TPJ, ACC, IFG Desire (Like vs. Hate) TPJ, ACC Overlap Orthogonal variation of beliefs and desires (Hartwright, Apperly & Hansen, 2012)

34. Belief (True vs. False) TPJ, ACC, IFG Desire (Like vs. Hate) TPJ, ACC Overlap Orthogonal variation of beliefs and desires (Hartwright, Apperly & Hansen, 2012) Notably no mPFC

35. Belief-desire task vs. ToM-localiser Belief OR Desire “ToM localiser” (False Belief – False Photo) Overlap Conjunction analysis between Belief-Desire and ToM Localiser

36. 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)

37. 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

38. 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

39. 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

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

41. Social abduction (Hartwright, Apperly & Hansen, subm.) Selective for D? TB vs. FB Green = D? vs. D-&D+ Green = D? vs. D-&D+&FB&TB

42. Case study 2 – Temporal coordination

43. Background: The neural basis of “theory of mind” Temporo-parietal junction Temporal pole Medial prefrontal cortex Left lateral view TPJ TP Medial view mPFC AnteriorPosterior PC TPJ TP mPFC lPFC Precuneus TPJ TP Anterior lPFC PC Lateral prefrontal cortex Right lateral view We don’t know how these regions work together e.g. Frith & Frith, 2003 Van Overwalle, 2009

44. Automatic perspective-taking? (Samson, Apperly, Braithwaite et al., 2010, JEP:HPP) You / He 2 2 Self / Other Consistent Self / Other Inconsistent Disc position varies 1,2, or 3 discs

45. You / He 2 2 Self / Other Consistent Self / Other Inconsistent Disc position varies 1,2, or 3 discs Automatic perspective-taking? (Samson, Apperly, Braithwaite et al., 2010, JEP:HPP)

46. Main effect of consistency Significant interaction RT (ms) Egocentric interference on explicit judgement of other Automatic perspective-taking? (Samson, Apperly, Braithwaite et al., 2010, JEP:HPP)

47. Main effect of consistency Significant interaction RT (ms) Altercentric interference =evidence of automatic calculation of perspective Automatic perspective-taking? (Samson, Apperly, Braithwaite et al., 2010, JEP:HPP)

48. Main effect of consistency Significant interaction RT (ms) Various follow-ups..... Altercentric interference = evidence of automatic calculation of perspective Automatic perspective-taking? (Samson, Apperly, Braithwaite et al., 2010, JEP:HPP)

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

50. * ns Automatic perspective-taking? (Samson, Apperly, Braithwaite et al., 2010, JEP:HPP) Only ever judge “self” – how many dots you can see

51. Automatic and controlled processes within a perspective-taking problem? Main effect of consistency Significant interaction RT (ms) Altercentric interference = indication of automatic perspective calculation CalculationSelectionResponse Self Yes Other

52. Automatic and controlled processes within a perspective-taking problem? Main effect of consistency Significant interaction RT (ms) Altercentric interference = indication of automatic perspective calculation Dual tasking CalculationSelectionResponse Self Yes Other

53. Cognitively effortful perspective selection Qureshi, Apperly & Samson (2010) Cognition. Altercentric interference is increased by dual tasking with an executive task

54. Background: The neural basis of “theory of mind” Temporo-parietal junction Temporal pole Medial prefrontal cortex Left lateral view TPJ TP Medial view mPFC AnteriorPosterior PC TPJ TP mPFC lPFC Precuneus TPJ TP Anterior lPFC PC Lateral prefrontal cortex Right lateral view e.g. Frith & Frith, 2003 Van Overwalle, 2009

55. Background: The neural basis of “theory of mind” Temporo-parietal junction Temporal pole Medial prefrontal cortex Left lateral view TPJ TP Medial view mPFC AnteriorPosterior PC TPJ TP mPFC lPFC Precuneus TPJ TP Anterior lPFC PC Lateral prefrontal cortex Right lateral view e.g. Frith & Frith, 2003 Van Overwalle, 2009

56. Background: The neural basis of “theory of mind” Temporo-parietal junction Temporal pole Medial prefrontal cortex Left lateral view TPJ TP Medial view mPFC AnteriorPosterior PC TPJ TP mPFC lPFC Precuneus TPJ TP Anterior lPFC PC Lateral prefrontal cortex Right lateral view e.g. Frith & Frith, 2003 Van Overwalle, 2009

57. Background: The neural basis of “theory of mind” Temporo-parietal junction Temporal pole Medial prefrontal cortex Left lateral view TPJ TP Medial view mPFC AnteriorPosterior PC TPJ TP mPFC lPFC Precuneus TPJ TP Anterior lPFC PC Lateral prefrontal cortex Right lateral view We don’t know how these regions work together e.g. Frith & Frith, 2003 Van Overwalle, 2009

58. 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.

59. 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

60. Perspective calculation: 450ms Self<Other latency over posterior scalp Confirmatory source analysis suggested Bilateral TPJ

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

62. 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

63. Conclusions Primacy 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 mPFC AnteriorPosterior PC TPJ TP mPFC lPFC Precuneus TPJ TP Anterior lPFC PC Lateral prefrontal cortex Right lateral view e.g. Frith & Frith, 2003 Van Overwalle, 2009

64. Conclusions Role for non-ToM “control network” in perspective selection Temporo-parietal junction Temporal pole Medial prefrontal cortex Left lateral view TPJ TP Medial view mPFC AnteriorPosterior PC TPJ TP mPFC lPFC Precuneus TPJ TP Anterior lPFC PC Lateral prefrontal cortex Right lateral view e.g. Frith & Frith, 2003 Van Overwalle, 2009

65. 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 mPFC PC

66. 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 mPFC PC What might we expect Mindreading to involve? The “ToM network”

67. 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 mPFC PC What might we expect Mindreading to involve? The “ToM network”

68. 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 mPFC PC ACC What might we expect Mindreading to involve? Cognitive control

69. 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 mPFC PC What might we expect Mindreading to involve? Cognitive control ACC

70. 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 mPFC PC What might we expect Mindreading to involve? Cognitive control ACC

71. 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 mPFC PC What might we expect Mindreading to involve? ACC

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

74. Main effect of Mental/non-mental in rTPJ Main effect of ambiguous/unambiguous in mPFC

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