1. Visual Attention Attention is the ability to select objects of interest from the surrounding environment A reliable measure of attention is eye movement during object (target) selection Early studies show that there are specific brain regions that are involved in the process of target selection

2. Visual Attention Attention is the ability to select objects of interest from the surrounding environment A reliable measure of attention is eye movement during object (target) selection Early studies show that there are specific brain regions that are involved in the process of target selection –Superior Colliculus

3. Visual Attention Attention is the ability to select objects of interest from the surrounding environment A reliable measure of attention is eye movement during object (target) selection Early studies show that there are specific brain regions that are involved in the process of target selection –Superior Colliculus –Frontal Eye Field

4. Visual Attention Attention is the ability to select objects of interest from the surrounding environment A reliable measure of attention is eye movement during object (target) selection Early studies show that there are specific brain regions that are involved in the process of target selection –Superior Colliculus –Frontal Eye Field –Extrastriate Areas (V1, V4, IT)

5. Superior Colliculus & Target Selection: Early Studies Cells in the superficial layers of the SC respond to light falling on a specific area of the retina Visual response occurs 50ms after stimulus onset even when the stimulus appears in the RF and the monkey maintains fixation

6. Superior Colliculus & Target Selection: Early Studies Enhanced visual response when the stimulus appears in the cell’s receptive field AND the monkey selects it as a target and saccades (overtly attends) to it. Latency of saccade is about 200ms Activity is greater for target + saccade than for target

7. Superior Colliculus & Target Selection: Early Studies No enhanced response when the monkey saccades to a stimulus outside the receptive field (or in total darkness)

8. Superior Colliculus & Target Selection: Early Studies No enhanced response when the monkey covertly attends to the stimulus in the receptive field At least this is what they found back then

9. Conclusions from Early Studies Early studies suggest that the superior colliculus is involved in the selection of targets for eye movements –Not just a visual response to stimulus –Not just a motor response to saccade Greatest level of activity is found when both a target is presented and a subsequent saccade is made to that target

10. Target Selection and Uncertainty Behavioral Task –Uncertainty period: array of possible targets presented –Selection period: one of the targets is dimmed –Removal of fixation signaled saccade initiation Manipulated uncertainty by varying the # of possible targets (distractors) Recorded activity in SC intermediate layer neurons

11. Visual Response Modulated by Uncertainty / # of Distractors Response during stimulus onset in the uncertainty period decreases as the # of distractors (uncertainty level) increases Response during selection and saccade initiation were not significantly different regardless of uncertainty

12. The Frontal Eye Field & Target Selection FEF cells respond to visual stimuli like the SC –Greater activity for spots of light in the receptive field –RF is larger: as big as a ¼ of an entire visual field

13. Decreased Activity with Increased Uncertainty / # of Distractors

14. Superficial Layers Intermediate and Deep Layers Extrastriate Cortex (e.g. V4, MT) Parietal Cortex (LIP) Frontal Eye Field Medio-Dorsal Thalamus Striate Cortex (V1) Retina Brainstem Saccade Generator SC Major Connections of the Superior Colliculus

15. Superficial Layers Intermediate and Deep Layers Extrastriate Cortex (e.g. V4, MT) Parietal Cortex (LIP) Frontal Eye Field Medio-Dorsal Thalamus Striate Cortex (V1) Retina Brainstem Saccade Generator SC Major Connections of the Superior Colliculus

16. Early Studies of Target Selection Activity in FEF Detection Task –Target presented alone –Goal: saccade to target Visual Search Task –Target with distractors –Goal: saccade to target Recorded neural activity in FEF at two time intervals: –100ms after stimulus onset –100ms before saccade

17. Selection-Dependent Visual Activity Enhanced response when the target is in the RF Greater response for target alone (detection) than with distractors (search) Suppressed activity when the target is near the RF only in search trials

18. Target Selection in FEF Neuron Detection Trials –Activity decreased as distance from RF increased Search Trials –Initial activity is the same, since a stimulus (distractor) is always present –Enhanced activity for target in RF –Suppressed activity for target near RF

19. Go / No-go Task Monkeys fixated on white spot Trigger signal: change of color –Go trial (solid square): shift gaze to the target – saccade –Nogo trial (asterisk): maintain fixation – no saccade Color change can occur before, during or after target onset

20. Neural Response for Go & No-go Trials

21. Evidence for Center-Surround Activity

22. FEF Connects to Extrastriate Cortex

23. Target Selection in V4

24. Visual Properties of IT Neurons Neurons respond selectively to complex objects –Input: V1 and V4 (feature detection) –Output: entorhinal cortex  hippocampus (memory) Large receptive fields that always include the fovea Retinotopic organization is absent Invariant Properties –Scale invariance: insensitive to changes in stimulus size –Translation invariance: insensitive to changes in position Columnar organization IT neurons are expected to be involved in more complex aspects of target selection

25. Experimental Design Good (square) and poor (triangle) stimuli were selected prior to task Sequence of Events –cue  delay  target –Saccade to target that matches the cue Block trial: used one cue for 20 – 30 trials Recorded activity in IT

26. IT Neurons Carry Target Information

27. Next Week’s Readings McPeek & Keller: effects of superior colliculus inactivation on target selection Bichot & Schall: responsiveness of FEF to target similarity and history Sheinberg & Logothetis: inferotemporal cortex and natural target selection of familiar objects