1. Post-test review session Tuesday Nov 22 4-5 in TH241

2. A Hard Problem Neuroscientists have deferred some of the difficulties of that problem by focusing on a subtly different one: What neural processes are distinctly associated with consciousness? –That is still a pretty hard problem! What are the neural correlates of consciousness (NCC)

3. Searching for the NCC What is needed is a situation in which a perceiver’s state can alternate between aware and unaware in ways that we can correlate with neural events One such situation is called Binocular Rivalry

4. Rivalrous Images A rivalrous image is one that switches between two mutually exclusive percepts

5. Binocular Rivalry What would happen if each eye receives incompatible input? Left EyeRight Eye

6. Binocular Rivalry What would happen if each eye receives incompatible input? The percept is not usually the amalgamation of the two images. Instead the images are often rivalrous. –Percept switches between the two possible images

7. Binocular Rivalry Rivalry does not entail suppression of one eye and dominance of another – it is based on parts of objects: Left EyeRight Eye Stimuli: Percept:Or

8. Binocular Rivalry Percept alternates randomly (not regularly) between dominance and suppression - on the order of seconds –What factors affect dominance and suppression? Time ->

9. Binocular Rivalry Percept alternates randomly (not regularly) between dominance and suppression - on the order of seconds –What factors affect dominance and suppression? –Several features tend to increase the time one image is dominant (visible) Higher contrast Brighter Motion

10. Binocular Rivalry Percept alternates randomly (not regularly) between dominance and suppression - on the order of seconds –What factors affect dominance and suppression? –Several features tend to increase the time one image is dominant (visible) Higher contrast Brighter Motion What are the neural correlates of Rivalry?

11. Neural Correlates of Rivalry What Brain areas “experience” rivalry? Clever fMRI experiment by Tong et al. (1998) –Exploit preferential responses by different regions –Present faces and buildings in alternation

12. Neural Correlates of Rivalry What Brain areas “experience” rivalry? Clever fMRI experiment by Tong et al. (1998) –Exploit preferential responses by different regions –Present faces to one eye and buildings to the other

13. Neural Correlates of Rivalry What Brain areas “experience” rivalry? Apparently activity in areas in ventral pathway correlates with awareness But at what stage is rivalry first manifested? For the answer we need to look to single-cell recording

14. Neural Correlates of Rivalry Neurophysiology of Rivalry –Monkey is trained to indicate which of two images it is perceiving (by pressing a lever) –One stimulus contains features to which a given recorded neuron is “tuned”, the other does not –What happens to neurons when their preferred stimulus is present but suppressed?

15. Neural Correlates of Rivalry The theory is that Neurons in the LGN mediate Rivalry

16. Neural Correlates of Rivalry The theory is that Neurons in the LGN mediate Rivalry NO – cells in LGN respond similarly regardless of whether their input is suppressed or dominant

17. Neural Correlates of Rivalry V1? V4? V5? YES – cells in primary and early extra-striate cortex respond with more action potentials when their preferred stimulus is dominant relative to when it is suppressed However, –Changes are small –Cells never stop firing altogether

18. Neural Correlates of Rivalry Inferior Temporal Cortex (Ventral Pathway)? YES – cells in IT are strongly correlated with percept

19. Neural Correlates of Rivalry Inferior Temporal Cortex (Ventral Pathway)? YES – cells in IT are strongly correlated with percept Why does area IT sound familiar to you?

20. Neural Mechanisms of Consciousness? So how far does that get us? Not all that far – we still don’t know what is the mechanism that causes consciousness But we do know that it is probably distributed rather than at one locus Thus the question is: what is special about the activity of networks of neurons that gives rise to consciousness? – that’s still a very hard problem

21. From Perception to Action And what’s in between?

22. A thought problem Mary the vision scientist has had a few drinks after a long day at a neuroscience conference. She boldly announces to the group at the table that Vision Science is the most important branch of neuroscience because “50% of the brain is visual cortex”. Charlie the motor control guy has also had a few drinks. “nonsense”, he exclaims, “50% of the brain is motor cortex so we must be equally important!” The rest of the group, comprised of auditory, memory and attention researchers start rolling up their sleeves and fixin for a fight, because that doesn’t leave any brain left for them to study.

23. A thought problem “Hold it!”, you shout, because you spot the problem immediately…

24. A thought problem “Hold it!”, you shout, because you spot the problem immediately…

25. The brain exhibits functional specialization, but the functions are complex and dynamic –E.g. catching a ball, driving a car, reading a book, running –Each example requires tight integration of perception and action

26. “motor systems” are perception systems as well

27. Some cells in basal ganglia have visual receptive fields that are locked to arm location rather than

28. The perception – action system

29. What are Mirror Neurons? Mirror neurons are cells that fire when a monkey (or person?) performs an action or when it views another animal performing that same action

30. What are Mirror Neurons? Located in conjunction with a frontoparietal action planning network: – posterior inferior frontal pre-motor areas that represent impending actions Many cells are “motor dominant”: they fire when monkey grasps or reaches, but are independent of visual input –inferior parietal sensory area that integrates somatosensory, visual and auditory signals Many cells are “visuo-dominant” or “visuomotor-dominant”: they fire when monkey sees a graspable object or a stimulus that could be interacted with –These regions are densely interconnected

31. What are Mirror Neurons? Mirror neurons are in regions immediately adjacent to these frontal and parietal areas Motor properties of mirror neurons are same as “non- mirror” neurons but… Sensory properties are different –These cells do not fire when monkey sees a graspable object –They do fire when monkey sees another monkey (or a person!) perform actions relative to objects

32. What can Mirror Neurons Represent? Some cells are precisely tuned to the specific actions (strictly congruent – about 1/3 of cells) –e.g. using two fingers to pick up an object Others are broadly tuned to any action that accomplishes the same goal (broadly congruent – about 2/3 of cells) –E.g. using any combination of hand and fingers to pick up an object

33. What can Mirror Neurons Represent? Mirror neurons represent abstract actions and goals –Cells will not fire when grasping is pantomimed –Cells will fire when grasping is real and visible –Cells will also fire when grasping happens behind an occluding screen, as long as the monkey has seen that there is an object to be grasped behind the screen!

34. What can Mirror Neurons Represent? Mirror neurons represent abstract actions and goals –Some mirror neurons will even fire in response to the sound of an action being performed e.g. the sound of breaking a peanut shell

35. What are mirror neurons for? Two theories: –Action imitation –Understanding actions of others

36. Mirror Neurons in Humans? The existence of mirror neurons in humans is inferred –We don’t record directly from human cortex!

37. Mirror Neurons in Humans? EEG and MEG evidence suggests a Mu desynchronization (9 – 12 Hz over premotor areas) when someone observes grasping TMS evidence shows that observed grasping decreases the threshold for triggering motor potentials for grasping muscles in the hand.

38. The putative Mirror Neuron System in humans Iacoboni et al (2006)

39. Mirror Neurons in Humans? fMRI studies have found evidence for two brain regions associated with imitative actions Task 1: watch action Task 2: perform action Task 3: imitate action

40. Mirror Neurons in Humans? Based on work with monkeys, it was presumed that mirror neuron areas would exhibit a specific pattern of BOLD response: Task 1 < Task 2 < Task 3 Two regions exhibited such a response

41. Mirror Neurons Dysfunction in Autism? Some evidence supports a theory that a dysfunction of the MNS underlies social isolation disorders such as autism spectrum disorder (ASD) –Some structural abnormalities revealed with anatomical MRI

42. Mirror Neurons Dysfunction in Autism? fMRI data shows that BOLD signal in MNS during viewing and imitating emotional expressions is negatively correlated with severity of Autism Spectrum Disorder

43. Mirror Neurons Dysfunction in Autism? Some evidence supports a theory that a dysfunction of the MNS underlies social isolation disorders such as autism spectrum disorder (ASD) –MEG data shows abnormal propagation of signals in the MNS when imitating lip movements in individuals with Asperger’s Syndrome