1. Social Cognition and the Mirror Neuron System of the Brain Jaime A. Pineda, Ph.D. Cognitive Neuroscience Laboratory COGS1 class

2. Motivating Questions How do our brains perceive the mental states of others despite their inaccessibility? How do we understand the actions, emotions and the intentions of others? Rationally? Intuitively? How do we understand first- and third-person experiences?

3. Classic Explanation Theory-Theory (argument from analogy; disembodied knowledge; visual hypothesis) Involves striate, extrastriate, inferotemporal lobe and superior temporal sulcus, among others

4. A Different Perspective Simulation Theory (Direct-matching hypothesis; embodied knowledge) Map visual information onto motor representations of the same action Mirroring systems bridges between perception and action that allow for simulation Mirror neurons EEG Mu rhythms

5. A Different Perspective Simulation Theory (Direct-matching hypothesis; embodied knowledge) Map visual information onto motor representations of the same action Mirroring systems bridges between perception and action that allow for simulation Mirror neurons EEG Mu rhythms

6. The Mirror Neuron System Iacoboni and Dapretto, Nature Reviews, 2006,7:942-951 Pineda, Beh & Brain Functions, 2008, 4, 47

7. Biological Motion Visual system's ability to recover object information from sparse input Gender Activity engaged in Emotional state

8. Biological Motion Perception: Monkeys Perret and colleagues (1989; 1990; 1994) Cells in superior temporal polysensory area (STPa) of the macaque temporal cortex appear sensitive to biological motion Oram & Perrett, J. Cog. Neurosci., 1994, 6(2), 99-116

9. Biological Motion Perception: Humans An area in the superior temporal sulcus (STS) in humans responds to biological motion Other areas do as well, including frontal cortex, SMA, insula, thalamus, amygdala Grossman et al. J. Cog. Neurosci., 2000, 12(5), 711-720

10. Brain Circuit for Social Perception (SP) Allison et al., Trends in Cog. Sci., 2000, 4, 267-272 SP is processing of information that results in the accurate analysis of the intentions of others STS involved in the processing of a variety of social signals

11. Mirror Neurons A specific class of neurons that discharge both when the monkey performs an action and when it observes a similar action done by another monkey or an experimenter Found in: area F5 (homolog of Broca’s area); 10-20% inferior parietal cortex (PF/7b) Activated by: Goal directed actions (reaching, grasping, holding) Observation of similar actions performed by “biological” agents Di Pellegrino et al., Exp. Brain Res., 1992, 91, 176-80

12. Rizzolatti et al., Cogn. Brain Res., 1996, 3:131-141 Mirror Neuron Activity

13. Perception-to-Action Mapping Selectivity Action Logically-Related (effector independent; 2X) Congruent (effector dependent) Perception

14. Understanding Intentions Umilta et al. Neuron, 2001, 32: 91-101 GraspingMimicking

15. Inferior parietal lobule Superior temporal sulcus Inferior frontal gyrus Sensorimotor cortex Mirror Neuron System

16. Response facilitation Mimicry Simulation Imitation learning Understanding actions Understanding intentions Empathy Theory of Mind Language Functional Significance

17. MNS activity No MNS Activity intentionality? anthropomorphism? biological realism? motivational significance? transitive/intransitive actions? generalizability? Characterizing the System learning? social relevance?

18. Mu Rhythm 8-13 Hz oscillation over sensorimotor cortex Normal Oscillation Self Action Observed Action

19. Frequency Analysis of Mu Rhythm PowerPower Frequency (8-13 Hz) (10-14 Hz)

20. Pineda et al., IEEE Trans. Rehab. Engr., 2000, 8(2): 219-222 Does Mu Suppression Reflect Mirror Activity ? Baseline Move Observe Imagine

21. Action Observation and Social Interaction To what degree do mu rhythms, like mirror neurons, reflect social interaction? Oberman et al., Social Cognitive and Affective Neuroscience, 2007, 2, 62-66

22. Measured mu power (2 min of EEG) in normals (n=20) ages 18-34 (mean=21.1, SD=3.40 ) under different observation conditions: Non-interacting Social Action - Spectator Social Action - Interactive Visual white noise Engaged in continuous performance task during observation Experimental Paradigm

23. Non-interactingSocial Action - Spectator Social Action - Interactive

24. Results

26. Autism Spectrum Disorder (ASD) Problems in the following domains: Social ability Language development Behavior

27. Common Characteristics Impairment in social play and imagination Impaired ability to initiate conversations with others Repetitive behaviors Impaired sustained attention Trouble Imitating others Difficulty interpreting actions and intentions of others Absence of empathy

28. Autism: A Dysfunctional Mirror System? No common underlying mechanism has been identified Deficits in imitation learning – Rogers and Pennington, 1991 Deficits in mirror neuron system - Williams et al., 2001

29. Hypothesis If mu rhythms reflect MNS activity and the capacity to understand actions as well as learn through imitation, then autistics should show differences in mu rhythms compared to controls Oberman et al., Cog. Brain Res. 2005, 24: 190-198

30. Experimental Paradigm Measured mu power (2 min of EEG) in normals (n=12) and autistics (n=10) under different conditions: Self-movement of hand Watching video of someone moving their hand Watching a video of a ball moving up and down Oberman et al., Brain Res Cogn Brain Res. 2005, 24(2):190-8.

31. Results

32. Oberman et al., Neuropsychologia, 2008 Jan 19 [Epub ahead of print]

33. Creating a Temporary “Autistic” Brain Do sensorimotor mu rhythms reflect downstream modulation from cells in premotor cortex? RATIONALE If mirror neurons in IFG are involved in the direct modulation of mu rhythms, then temporary inhibition of these neurons should prevent suppression of mu rhythms and cause “autistic-like” behaviors.

34. Method Measured EEG mu power in typically developing adults (n=8) under different conditions before and after IFG stimulation Observation of movement (4 videos) Simple (hand movements) and complex (social interactions) Baron-Cohen’s Eyes Task Emotion and gender discrimination 1 Hz rTMS (5 min at ~ 40-50% MEP threshold) targeted at left IFG

35. Eyes Task

36. Results Reaction Time Accuracy

37. Results

38. Frontoparietal areas in an ASD brain may be underconnected If we change the dynamics of the sensorimotor mu oscillations, And these oscillations are functionally linked to the MNS network (IFG, IPL, STS), Then we may change functional connectivity and recover MNS engagement, Leading to positive changes. IPL STS SM Cortex IFG Plasticity Induced Rehabilitation Training

39. Reversing Social Deficits in Autism? Training 30 min x 3/week x 10 weeks HF ASD: 7-17 yr olds; n=20 Experimental/Control groups Mu activity above threshold (E) EMG activity below threshold (E/C) Pineda et al., Research in ASD, 2008

40. Pre/Post Assessments Verification of diagnosis (IQ, ADI, ADOS) Quantitative EEG (QEEG) Test of Variable Attention (TOVA) Imitation ability (Apraxia imitation test) Mu suppression index (MSI) Autism Treatment Evaluation Checklist (ATEC) Neuroimaging (fMRI, fcMRI)

41. Facial Emotions (n-back)Eyes Emotion Task - ToM Physical Causation - ToM Mental Attribution- ToM Pre/Post Assessments

42. ASD TD Behavioral Performance

43. Mu Suppression Index

44. EEG Coherence

45. Sustained Attention Reduction trend in ADHD score for experimental group

46. Autism Treatment Evaluation Checklist

47. Functional connectivity Diffusion Tensor Imaging structural functional Future: Neuroimaging Techniques

48. A Fundamental Organizational Feature of the Brain? “Understanding others as intentional agents may be grounded in the relational nature of our interactions with the world” Beyond understanding actions emotions: the root of empathy? sounds and other senses language Other problems in “mirroring” Aberrant imitation learning: addiction?

49. What Is It Like To Be…? Thomas Nagel, The Philosophical Review 83 (1974). Can aspects of subjective experience be reduced to brain activity?

50. Collaborators and Students Vilayanur Ramachandran Lindsay Oberman Eric Altschuler Andrey Vankov Bill Skinner Chulie Ulloa Brendan Allison Ed Hubbard Joe McCleery Erin Hecht David Brang Scott Carey Adrienne Moore Rajiv Rao Chris Robinson Hanie Elfenbein Alex Bressler Steven Thurman Jena Davis Dong Suk Christa Futagaki Judith Kaye Lee Edwards Ralph-Axel Mueller Brandon Keehn Oriana Clark Jia-Min Bai Derrick Asher Dane Chambers Matt Earhardt Heather Pelton Alicia Trigerio Albert Ayala Stephen Johnson Steve Gilmore Nick Pojman Kelly Head