1. Read this article for Friday Oct 21! Trends in Neuroscience (2000) 23, 571 - 579 Hint #1: there are at least 3 ways of getting this article Hint #2: none of them are “wait till Matt sends it to me”

2. Visual Pathways Retina has distinct layers

3. Visual Pathways Retina has distinct layers Amacrine and bipolar cells perform “early” processing –Peripheral retina features convergence of receptors onto ganglion cells –Foveal retina features divergence –Why!?

4. Visual Pathways Retina has distinct layers Amacrine and bipolar cells perform “early” processing –lateral inhibition at middle layer leads to centre-surround receptive fields – first step in shaping “tuning properties” of higher-level neurons

5. Visual Pathways Retina has distinct layers –Ganglion cells project mainly to Lateral Geniculate Nucleus (LGN) of the thalamus –two kinds of ganglion cells: Magnocellular and Parvocellular Magno is myelinated Parvo is not myelinated In what way does this matter? –visual information is already being shunted through functionally distinct pathways

6. Visual Pathways visual hemifields project contralaterally –exception: bilateral representation of fovea! Optic nerve splits at optic chiasm about 90 % of fibers project to cortex via LGN about 10 % project through superior colliculus and pulvinar –but that’s still a lot of fibers! Note: this will be important when we talk about visuospatial attention

7. Visual Pathways Lateral Geniculate Nucleus maintains segregation: –of M and P cells (mango and parvo) –of left and right eyes P cells project to layers 3 - 6 M cells project to layers 1 and 2

8. Visual Pathways Cortical regions vary in their anatomical connections and their functional specialization Number should be thought of very loosely as “consecutive” as in a processing hierarchy –But note that this view is outdated as we’ll discover in reading the article by Lamme

9. Visual Pathways Primary visual cortex receives input from LGN –also known as “striate” because it appears striped when labeled with some dyes –also known as V1 –also known as Brodmann Area 17

10. Visual Pathways W. W. Norton Primary cortex maintains distinct pathways – functional segregation M and P pathways synapse in different layers Ascending (i.e. feed-forward) projections synapse in middle layers Descending (i.e. feed-back) projections synapse in superfical and deep layers

11. Visual Pathways Visual scene is represented: –Retinotopically thus… –spatiotopically = Fovea Tootell R B H et al. PNAS 1998;95:811-817

12. How does the visual system represent visual information? How does the visual system represent features of scenes? Vision is analytical - the system breaks down the scene into distinct kinds of features and represents them in functionally segregated pathways

13. Visual Neuron Responses The notion of a receptive field is fundamental in vision science –A neuron’s receptive field is the region in space in which a stimulus will evoke a response from that neuron –Receptive field properties vary widely across visual neurons and are never just “ON” or “OFF” –Unit recordings in LGN reveal a centre/surround receptive field

14. Visual Neuron Responses Unit recordings in LGN reveal a centre/surround receptive field many arrangements exist, but the “classical” RF has an excitatory centre and an inhibitory surround these receptive fields tend to be circular - they are not orientation specific How could the outputs of such cells be transformed into a cell with orientation specificity?

15. Visual Neuron Responses LGN cells converge on “simple” cells in V1 imparting orientation (and location) specificity

16. Visual Neuron Responses LGN cells converge on “simple” cells in V1 imparting orientation (and location) specificity Again, information is physically seperated into a “map”

17. Visual Neuron Responses LGN cells converge on simple cells in V1 imparting orientation specificity Thus we begin to see how a simple representation – orientations of lines - can be maintained in the visual system –increase in spike rate of specific neurons indicates presence of a line with a specific orientation at a specific location on the retina –Reality is that spike rate probably is only one part of the story: information is coded in many ways e.g. Relative timing Graded potentials

18. The Role of “Extrastriate” Areas Different visual cortex regions contain cells with different tuning properties

19. The Role of “Extrastriate” Areas Consider two plausible models: 1.System is hierarchical: –each area performs some elaboration on the input it is given and then passes on that elaboration as input to the next “higher” area 2.System is analytic and parallel: –different areas elaborate on different features of the input

20. The Role of “Extrastriate” Areas Functional imaging (PET) investigations of motion and colour selective visual cortical areas Zeki et al. Subtractive Logic –stimulus alternates between two scenes that differ only in the feature of interest (i.e. colour, motion, etc.)

21. The Role of “Extrastriate” Areas Identifying colour sensitive regions Subtract Voxel intensities during these scans… …from voxel intensities during these scans …etc. Time ->

22. The Role of “Extrastriate” Areas result –voxels are identified that are preferentially selective for colour –these tend to cluster in anterior/inferior occipital lobe

23. The Role of “Extrastriate” Areas similar logic was used to find motion-selective areas Subtract Voxel intensities during these scans… …from voxel intensities during these scans …etc. Time -> MOVING STATIONARY MOVING STATIONARY

24. The Role of “Extrastriate” Areas result –voxels are identified that are preferentially selective for motion –these tend to cluster in superior/dorsal occipital lobe near TemporoParietal Junction –Akin to Human V5

25. The Role of “Extrastriate” Areas Thus PET studies doubly-dissociate colour and motion sensitive regions

26. The Role of “Extrastriate” Areas V4 and V5 are doubly-dissociated in lesion literature:

27. The Role of “Extrastriate” Areas V4 and V5 are doubly-dissociated in lesion literature: –achromatopsia (color blindness): there are many forms of color blindness cortical achromatopsia arises from lesions in the area of V4 singly dissociable from motion perception deficit - patients with V4 lesions have other visual problems, but motion perception is substantially spared

28. The Role of “Extrastriate” Areas V4 and V5 are doubly-dissociated in lesion literature: –akinetopsia (motion blindness): bilateral lesions to area V5 (extremely rare) severe impairment in judging direction and velocity of motion - especially with fast-moving stimuli visual world appeared to progress in still frames similar effects occur when M-cell layers in LGN are lesioned in monkeys

29. Visual Neuron Responses Edges are important because they are the boundaries between objects and the background or objects and other objects

30. Visual Neuron Responses This conceptualization of the visual system was “static” - it did not take into account the possibility that visual cells might change their response selectivity over time –Logic went like this: if the cell is firing, its preferred line/edge must be present and… –if the preferred line/edge is present, the cell must be firing We will encounter examples in which these don’t apply! Representing boundaries must be more complicated than simple edge detection!

31. Visual Neuron Responses Boundaries between objects can be defined by color rather than brightness

32. Visual Neuron Responses Boundaries between objects can be defined by texture

33. Visual Neuron Responses Boundaries between objects can be defined by motion and depth cues