Attention March 4, 2009

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1 Attention March 4, 29 John Maunsell Mitchell, JF, Sundberg, KA, Reynolds, JH (27) Differential attention-dependent response modulation across cell classes in macaque visual area V4. Neuron, 55:

4 Moore et al.

6 Spatial attention task receptive field 1. fixation point fraction of probes detected. attended location unattended location attend to other stimulus attend to receptive field stimulus neuronal response (spikes/s) time (ms) 1 time (ms) 1

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9 Task difficulty modulates neuronal responses in Area V4 Spitzer, Moran & Desimone 1988

12 Detecting orientation change Boudreau et al. (26) fixation point receptive field easy mode (9 ) difficult mode (~1 )

13 Task difficulty is determined by the size of orientation change Boudreau et al. (26) easy difficult 9 ~1

14 Task difficulty is determined by the size of orientation change Boudreau et al. (26) easy difficult 9 ~1 probe ~2 probe ~2

15 Behavioral performance Boudreau et al. (26) 1. (n=872) performance on probes (proportion correct).8.6 easy mode.4 difficult mode.2. attended unattended

16 Example V4 neuron response to preferred orientation Boudreau et al. (26) 3 response (spikes/s) 3 difficult mode attended unattended 25 time within stimulus cycle (ms) 3 easy mode 25 time within stimulus cycle (ms) neuronal response (spikes/s) 2 easy mode 1 difficult mode attended unattended

17 Distribution of attentional modulation in V4 Boudreau et al. (26) easy mode difficult mode 4 median = 7% 4 median = 24% number of neurons attentional modulation (attended / unattended) attentional modulation (attended / unattended)

18 !"##$%&'())* #%:;#3"2 #"&<3=#.##$2#3"2,-$%./$ %$1"2$ '134$5+ 6 9*66 6 * >3?$&'?+

19 Average attention modulation of single unit responses at different cortical levels Cook & Maunsell (22) 1% average response enhancement by attention 5% % V1 V4 MT VIP MST 7a cortical hierarchy levels

20 !"#$%&'%(&)*+,-"&.%/,-",+%,--$.-"&.%&'%'123%/"4.,+%".%)"''$5$.-%6"/*,+%,5$,/% (&)*+,-"&.% ".)$F% BGH% BGB% DE% DA% DL% DLM% DK% 1I% IJ7%

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22 Modulation of V1 fmri BOLD signal by attention Somers et al A 1 cm p V1+ V1-

23 Recording from human visual cortex Yoshor et al voltage (mv)!1! time (ms)

24 Responsive and unresponsive locations Yoshor et al elevation azimuth azimuth -8-4 azimuth voltage (µv) voltage (µv) 2 time (ms) 2 time (ms)

25 Spatial attention task Yoshor et al fixation elevation -4 stimuli response period -8-4 azimuth 4 time

26 Effect of attention on human LFP Yoshor et al percent correct valid cue invalid cue voltage (µv) orientation change time (ms)

27 stronger response response stronger response & increased selectivity orientation

28 Change in average V4 orientation tuning by attention 1. attend in attend out n = 262 V4 sample response.5 amplitude:.45.6 width: orientation relative to preferred 9

29 Attention does not sharpen direction tuning in MT Treue & Martinez-Trujillo 1999 pattern A pattern B 8 attend to pattern B response (spikes/s) attend to pattern A stimulus direction pattern B

30 Average MT contrast response functions (n = 56) 6 average rate of firing (spikes/s) contrast (%)

31 Attention does not sharpen direction tuning in MT Treue & Martinez-Trujillo 1999 pattern A pattern B 8 attend to pattern B response (spikes/s) attend to pattern A stimulus direction pattern B

32 Attention can sharpen the tuning of a population response Martinez-Trujillo & Treue 24

33 Orientation tuning measured with different contrasts Cat V1 Sclar & Freeman % contrast response (spikes/s) 4% 2% 1% orientation

34 Change in average V4 orientation tuning by attention McAdams & Maunsell (1999) 1. attend in attend out n = 262 V4 sample response.5 amplitude:.45.6 width: orientation relative to preferred 9

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37 Variance of cat V1 neuron response Dean vary: spatial frequency contrast response variance (spikes 2 ) response mean (spikes)

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45 MT responses with attention to one of two receptive field stimuli Treue & Maunsell, 1996 fixation point stim. 1 attend to stimulus 1 attend to stimulus 2 stim response (spikes/s) 365 time (ms) 365 time (ms)

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47 Shifting attention between two stimuli inside the receptive field of a neuron in MT location fixation point location 1 location 2 stimulus receptive field location 2 blank blank null intermediate preferred attend location attend location 1 attend location 2 null location 1 stimulus intermediate preferred

48 Responses of an MT neuron to different stimulus pairings and attentional states location 2 stimulus blank null intermediate preferred blank null location 1 stimulus intermediate preferred attend away attend location 1 attend location 2 response 19 2 time (ms)

49 MT population average responses to different stimulus pairings and attentional states location 2 stimulus blank null intermediate preferred blank null location 1 stimulus intermediate preferred attend away attend location 1 attend location 2 response time (ms)

50 MT neuron 126 sp/s 1 s 5

51 Normalization of neuronal responses preferred stimulus rate of firing (spikes/s) 1 25 time (ms) 1 rate of firing (spikes/s)

52 Normalization of neuronal responses non-preferred stimulus rate of firing (spikes/s) 1 25 time (ms) 1 rate of firing (spikes/s)

53 Normalization of neuronal responses preferred stimulus non-preferred stimulus rate of firing (spikes/s) 1 25 time (ms) 1 rate of firing (spikes/s)

54 Normalization of neuronal responses Heeger et al nearby cells preferred stimulus rate of firing linear operator normalization output nonlinearity

55 Normalization of neuronal responses Heeger et al nearby cells preferred stimulus non-preferred stimulus linear operator output nonlinearity rate of firing nearby cells

56 Attention Normalization Model Lee & Maunsell (29) divisive normalization + excitatory + inputs MT neuron firing rate R 1,2 "!!"#!! #! #! " # # attended: $ " " # # # #! " # # $! $ attended! I 1 I 2 #! % " $! %% $! & '("! % Normalization pool N 1 N 2 ignored: #! % " $! %% $! & '(" % preferred null

57 MT population average responses to different stimulus pairings and attentional states location 2 stimulus blank null intermediate preferred blank null location 1 stimulus intermediate preferred attend away attend location 1 attend location 2 response time (ms)

58 Responses of an MT neuron and fit of normalization model location 2 stimulus blank null intermediate preferred blank null location 1 stimulus intermediate 93 preferred out loc 1 loc 2 attention to out loc 1 loc 2 attention to out loc 1 loc 2 attention to response out loc 1 loc 2 attention to

59 Two stimuli in the receptive field with different contrasts cued location cued location non-preferred 5% non-preferred 1% preferred 1% preferred 5%

60 Effect of contrast offset on the response functions of an MT neuron attention fixed relative contrast varied null 5% preferred 1% 135 firing rate (spikes/s) preferred stimulus contrast (%)

61 Effect of contrast offset on the response functions of an MT neuron attention fixed relative contrast varied null 5% preferred 1% 135 preferred 1% null 5% firing rate (spikes/s) preferred stimulus contrast (%)

62 Effect of contrast offset on the response functions of an MT neuron attention fixed relative contrast varied null 5% preferred 1% 135 preferred 1% null 5% firing rate (spikes/s) preferred 5% null 1% preferred stimulus contrast (%)

63 Contrast response functions of an MT neuron with strong modulation attention fixed relative contrast varied attention varied relative contrast fixed null 5% null preferred 1% preferred 135 firing rate (spikes/s) preferred stimulus contrast (%) contrast (%)

64 Contrast response functions of an MT neuron with weak modulation attention fixed relative contrast varied attention varied relative contrast fixed null 5% null preferred 1% preferred 55 firing rate (spikes/s) preferred stimulus contrast (%) contrast (%)

65 Modulation by attention is correlated to modulation by stimulus interactions 1 r =.76 p <.1 95% CIs modulation by stimulus interactions (attention fixed relative contrast varied) modulation by attention (attention varied relative contrast fixed)

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Key questions about attention

Key questions about attention How does attention affect behavioral performance? Can attention affect the appearance of things? How does spatial and feature-based attention affect neuronal responses in