1. Covert Attention Mariel Velez 4-28-2005

2. What is attention? Attention is the ability to select objects of interest from the surrounding environment Involuntary vs Voluntary Spatial vs Object Overt vs. Covert Attending to a stimulus enhances neural response to that stimulus Salience: Represents how important a visual signal is: adds weights to incoming signals according to some feature

3. Ability to attend to a stimulus without shifting one’s gaze towards it Direct gaze may be interpreted as hostile Covert Attention

4. Frontal Eye Fields Posterier Parietal Cortex (LIP) SC V4 1 2 3 4 FEF, SC saccade-only enhancement

5. LIP, FEF, SC Bisley JW and Goldberg ME. Neuronal activity in the lateral intraparietal area and spatial attention. Science 299:81-86, 2003 Moore T, Armstrong KM. Selective gating of visual signals by microstimulation of frontal cortex. Nature 421:370-3, 2003 Cavanaugh, J and Wurtz, R.H. Subcortical Modulation of Attention Counters Change Blindness. JofNeuroscience 24(50): 11236- 11243, 2004

6. Effects of attention on PPC Mountcastle VS Goldberg and Friends

7. Lynch, Mountcastle, Talbot and Yin-1977 MOTOR COMMAND HYPOTHESIS-presaccadic burst specific to saccade Recording from Area 7 (Posterior Parietal Cortex “Saccade Neurons”- presaccadic burst only when monkey makes a saccade— NOT activated by visual stimulus

8. Bushnell, Goldberg, Robinson 1981 “Impossible to determine whether the relationship of neuronal response to eye movements was specific to that movement or more related to the attentional mechanisms that are associated with eye movement” Need to dissociate the oculomotor process with the attention of the stimulus---- COVERT ATTENTION ATTENTION-Enhancements of presaccadic activity in the absence of saccades

9. Goldberg Task What is the relationship between LIP activity and enhanced behavioral performance during attention? Correlate firing of LIP with performance

10. Effects of attention on visual cortex

11. Moran and Desimone 1985 Delayed Match to Sample Task Attention filters out irrelevant stimuli

12. Attentional effects all over visual cortex-MT, MST (Treue and Maunsell J Neurosci. 1999 ) When one of the receptive field stimuli was the attended dot, the response of the neuron was strong whenever that dot moved in the preferred direction

13. McAdams and Maunsell (1999) J Neurosci. 19:431-441. Monkey attends to receptive field stimulus Receptive Field (RF) of a V4 neuron RF stimulus Monkey attends elsewhere Attention modulates V4 tuning

14. V4 But attentional signals can represent motor preparation (intention ) AND visual selection (attention) Information about visual targets guides the saccade Fovea’s landing point along the bar could be predicted by the degree to which V4 cells coded that bar prior to the saccade (Moore 1999)

15. Role of oculomotor mechanisms in spatial attention- FEF, SC

16. FEF Stimulation evokes saccades-Amplitude and direction of saccades are organized retinotopically Goldberg-no covert attn effects. Related to saccades specifically Reciprocally connected to lots of posterior visual areas including V4 Should be able to drive spatial attn by perturbing oculomotor signals

17. Moore 2003 How is FEF modulating individual V4 neurons? Is the FEF an oculomotor salience map?

18. SC: target selection vs attention SC presaccadic activity—gateway to the Brainstem Saccade Generator McPeek and Keller (2004) SC inactivation causes defects in target selection by reducing behavioral salience. SC is providing some info to visual cortical areas: Can SC microstimulation affect attention?

19. Wurtz’ Change Blind Task Replace visual cue with SC microstimulation to see if this counters change blindness Change blindness-”failure to see large changes in a visual scene that occur simultaneously with a global transient (ie blanks between visual scenes)” Cue to the area of visual change counters change blindness

20. Change Blind Task ftp://lsr-ftp.nei.nih.gov/web/jc/cb_demo.htm

21. Salience map Represents how important a visual signal is: adds weights to incoming signals according to some feature “ Activation of a particular subset of the map would strengthen the representation of whatever stimulus is positioned at the corresponding point in space, while failing to alter its identity” Where is (are) the salience map (s)? What other features are in a salience map? Is there a corresponding “not salient map” ? Are there multiple salience maps? How would these multiple maps interact? Top down vs bottom up?

22. Red-Bottom up Waldo-Top down

23. LIP, FEF, SC Bisley JW and Goldberg ME. Neuronal activity in the lateral intraparietal area and spatial attention. Science 299:81-86, 2003 Moore T, Armstrong KM. Selective gating of visual signals by microstimulation of frontal cortex. Nature 421:370-3, 2003 Cavanaugh, J and Wurtz, R.H. Subcortical Modulation of Attention Counters Change Blindness. JofNeuroscience 24(50): 11236- 11243, 2004

24. Covert Attention-Early Psychophysical Studies Premotor Theory—(Rizzolatti et al. 1983, 1987) Subjects instructed to hit button as soon as the stimulus appeared RT increased when stimulus is presented in a location different than the attended one. An even larger increase in RT occurs when stimulus appears in non-attended location in the opposite hemifield Premotor Theory-Motor Program controls covert orienting: distance and direction changes modify the program which increases the RT