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Attentive tracking of multiple objects by humans and monkeys 30 Summary Can Monkeys Mentally Track Multiple Stimuli? 1 2 3 4 0 20 40 60 80 100 1 2 3 0 20 40 60 80 100 1 2 3 0 20 40 60 80 100 Monkey M % C o r r e c t Items Tracked Monkey B Items Tracked Human (N=5) Items Tracked 4 Items 4 Items 5 Items 5 Items 6 Items 6 Items 8 Items 8 Items Chance Chance 1) Monkeys can track 2 and humans can track 3-4 items. 2) Humans discriminate form changes better on targets. 3) In V4 tracked stimuli elicit higher firing rates. 4) This response increase is not measurably diminished when a second stimulus is tracked. Jude F. Mitchell, Kristy A. Sundberg & John H. Reynolds 18 of 29 neurons show significantly higher firing for targets in either 1 of 4 or in 2 of 4 tracking tasks. * Monkeys made saccades to all targets and to no distracters to receive reward. * Trajectories were identical to the monkey task. * Rings were replaced with figure 8’s. * At the motion pause, all items changed from 8’s to E’s or 3’s, except one that changed to either a 2 or 5. * Subjects tracked the targets and also reported by key-press whether a 2 or 5 was present. Attention Task: V4 responses are enhanced for tracked items. The strength of modulation is similar for tracking 1 or 2 targets. References 1. Pylyshyn ZW and Storm RW (1988) Tracking multiple independent targets: evidence for a parallel tracking mechanism. Spat Vis. 3(3):179-97. 2. Sears CR and Pylyshyn ZW (2000) Multiple object tracking and attentional processing. Can J Exp Psychol. 54(1):1-14. * Systems Neurobiology Laboratory, The Salk Institute for Biological Studies. Funding provided by NIH Training Grant in Cognitive Neuroscience (J.M.), NSF Graduate Research Fellowship (K.S.), and NEI grant 1R01EY13802 (J.R.). Task: while maintaining fixation, track all cued stimuli. 4-8 identcal stimuli appear (200ms) 1-4 targets cued by luminance increment (200ms) Stimuli move to pause position (750 ms) Stimuli pause (300ms) Stimuli move to final position (750 ms) Cue off (200ms) Fixation point disappears: saccade To ALL targets and NO distracters to earn reward Yes. Monkeys can track at least 2 stimuli Are tracked stimuli attended? 20 40 60 80 100 Human (N=5) Target % Correct 1 of 4 2 of 4 1 of 5 2 of 5 1 of 6 2 of 6 3 of 6 Items Tracked 1 of 4 2 of 4 1 of 5 2 of 5 1 of 6 2 of 6 3 of 6 Items Tracked Distracter Yes. Discrimination is better for tracked stimuli. * Are V4 Neurons Modulated? * The monkey attentively tracked targets far from the receptive field while oriented, colored gabors appeared throughout the visual field. The neuron’s spatial receptive field, color, and orientation tuning were computed on the basis of responses to the gabors. . Step 1) Mapping a V4 Unit’s Receptive Field 4 identical stimuli appear (200ms) 1 targets cued by luminance increment (100ms) Stimuli move to pause position (750 ms) Stimuli pause (1000ms) Stimuli move to final position (750 ms) Cue off (100ms) Fixation point off: saccade to the target Tracking Away from RF Flashing Gabors + RF RF RF RF RF RF RF RF RF RF RF RF RF RF 4 identical stimuli appear (200ms) 1 targets cued by luminance increment (100ms) Stimuli move to pause position (750 ms) Stimuli pause (1000ms) Stimuli move to final position (750 ms) Cue off (100ms) Fixation point off: saccade to the target Tracking Away from RF Flashing Gabors + RF RF RF RF RF RF RF RF RF RF RF RF RF RF RF RF RF RF RF RF RF RF RF RF RF RF RF RF 0 100 0 100 0 100 0 100 0 100 0 100 0 100 2 4 6 8 10 12 2 4 6 8 10 12 2 4 6 8 10 12 2 4 6 8 10 12 2 4 6 8 10 12 2 4 6 8 10 12 0 100 Response to Flashed Gabor Image In x,y Space broken down by Orientation # of Repeats per x,y position 0-33 ms 33-67 ms 67-100 ms 100-133 ms 133-167 ms 167-200 ms Response Window TIME x y + x y + +1.5 -8.75 -6.25 +11.25 Fixation x y + +1.5 -8.75 -6.25 +11.25 Fixation Example Neuron: 0 100 200 0 100 200 0 100 200 0 100 200 0 100 200 2 4 6 8 10 12 2 4 6 8 10 12 2 4 6 8 10 12 2 4 6 8 10 12 2 4 6 8 10 12 2 4 6 8 10 12 0 100 200 # of Repeats per x,y position 0-33 ms 33-67 ms 67-100 ms 100-133 ms 133-167 ms 167-200 ms TIME y x Response to Flashed Gabor Image In x,y Space broken down by Color + Joint Color/Orientation Tuning for Example Cell: 2 4 6 8 2 4 6 8 2 4 6 8 2 4 6 8 2 4 6 8 2 4 6 8 0 500 1000 5 10 15 20 5 10 15 20 5 10 15 20 5 10 15 20 5 10 15 20 5 10 15 20 0 50 100 0-33 ms 33-67 ms 67-100 ms 100-133 ms 133-167 ms 167-200 ms Response Window TIME Spatial Response Orientation & Color Response y x # of Repeats Orientation Color X – stat. sig. > baseline O – stat. sig. < baseline 4 identical stimuli appear (200ms) 1 or 2 targets cued by luminance increment (100ms) Stimuli move to pause position (750 ms) Stimuli Pause (1000 ms) Stimuli move to final position (750 ms) Cue off (100ms) Fixation point off: saccade to ALL targets RF RF RF RF RF RF RF RF RF RF RF RF RF RF RF 20 40 60 80 100 Percent Correct Performance over Recording Sessions 1 of 4 Tracking 2 of 4 Tracking Step 2) Preferred Stimuli Enter and Pause in the Unit’s Receptive Field During Tracking We recorded 29 cells from Monkey M: 1000 1500 2000 2500 3000 10 20 30 40 50 60 70 Time (ms) Normalized Response Mean Response (N=29, All Cells) Pause Resume Attended Unattended Attended Unattended 1000 1500 2000 2500 3000 10 20 30 40 50 60 70 Time (ms) Normalized Response Pause Resume Attended Unattended Attended Unattended 0 10 20 30 40 50 Peak Normalized Response Modulation = 21.4% 0 10 20 30 40 50 Peak Normalized Response Modulation = 29.0% 1 of 4 Tracking 2 of 4 Tracking AI Index 1 of 4 Tracking AI Index 2 of 4 Tracking Relative Attentional Modulation for 1 or 2 Target Tracking -0.2 -0.1 0 0.1 0.2 0.3 -0.2 -0.1 0 0.1 0.2 0.3 AI = Attended - Distracter Attended + Distracter Corr. = 0.76 Neuron Shown Above * Stat. Sig. o Non-Sig. * Stat. Sig. o Non-Sig. There were no significant differences in the strength of modulation for the tracking of a single or two stimuli. Pause Period Stimuli pause (75 ms) Stimuli pause (75 ms) Form Change (150 ms) Example Cell: Unit C20.1: 1 of 4 Tracking 500 1000 1500 2000 2500 3000 20 40 60 80 100 120 140 500 1000 1500 2000 2500 3000 20 40 60 80 100 120 140 Appear Move Pause Resume End Trial Number 0 500 1000 1500 2000 2500 3000 3500 0 5 10 15 20 25 30 Firing Rate (Hz) Time (ms) 0 5 10 15 20 Firing Rate (Hz) Target Distracter 0 5 10 15 20 g ( 500 1000 1500 2000 2500 3000 50 100 150 200 500 1000 1500 2000 2500 3000 50 100 150 200 Appear Move Pause Resume End 0 500 1000 1500 2000 2500 3000 3500 0 5 10 15 20 25 30 Trial Number Firing Rate (Hz) Time (ms) Firin Rate Hz) Target Distracter Unit C20.1: 2 of 4 Tracking Gabor duration 33 msec., One gabor appeared every 8 msec., Resulting in 4 gabors being present at any point in time. Response Window Target Distracter Target Distracter

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Attentive tracking of multiple objects by humans and monkeys

30

Summary

Can Monkeys Mentally Track Multiple Stimuli?

1 2 3 40

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100

1 2 30

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1 2 30

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Monkey M

%Correct

Items Tracked

Monkey B

Items Tracked

Human (N=5)

Items Tracked

4 Items4 Items

5 Items5 Items

6 Items6 Items

8 Items8 Items

ChanceChance

1) Monkeys can track 2 and humans can track 3-4 items.2) Humans discriminate form changes better on targets.3) In V4 tracked stimuli elicit higher firing rates.4) This response increase is not measurably diminished when a second stimulus is tracked.

Jude F. Mitchell, Kristy A. Sundberg & John H. Reynolds

18 of 29 neurons show significantly higher firing for targets in either 1 of 4 or in 2 of 4 tracking tasks.

* Monkeys made saccades to all targets and to no distracters to receive reward.

* Trajectories were identical to the monkey task. * Rings were replaced with figure 8’s.* At the motion pause, all items changed from 8’s to E’s or 3’s, except one that changed to either a 2 or 5. * Subjects tracked the targets and also reported by key-press whether a 2 or 5 was present.

Attention Task:

V4 responses are enhanced for tracked items. The strength of modulation issimilar for tracking 1 or 2 targets.

References1. Pylyshyn ZW and Storm RW (1988) Tracking multiple independent targets: evidence for a parallel tracking mechanism. Spat Vis. 3(3):179-97.2. Sears CR and Pylyshyn ZW (2000) Multiple object tracking and attentional processing. Can J Exp Psychol. 54(1):1-14.

* Systems Neurobiology Laboratory, The Salk Institute for Biological Studies. Funding provided by NIH Training Grant in Cognitive Neuroscience (J.M.), NSF Graduate Research Fellowship (K.S.), and NEI grant 1R01EY13802 (J.R.).

Task: while maintaining fixation, track all cued stimuli.

4-8 identcal stimuli appear

(200ms)

1-4 targets cued by luminance increment

(200ms)

Stimuli move to pause position

(750 ms)

Stimuli pause(300ms)

Stimuli move tofinal position

(750 ms)Cue off(200ms)

Fixation point disappears: saccade

To ALL targets and NO distracters to earn reward

Yes.Monkeys can track at least 2 stimuli

Are tracked stimuli attended?

20

40

60

80

100

Human (N=5)

Target

% C

orr

ec

t

1 of 4 2 of 4 1 of 5 2 of 5 1 of 6 2 of 6 3 of 6

Items Tracked

1 of 4 2 of 4 1 of 5 2 of 5 1 of 6 2 of 6 3 of 6

Items Tracked

Distracter

Yes.Discriminationis betterfor trackedstimuli.

*

Are V4 Neurons Modulated?

* The monkey attentively tracked targets far from the receptive field while oriented, colored gabors appeared throughout the visual field. The neuron’s spatial receptive field,color, and orientation tuning were computed on the basis of responses to the gabors..

Step 1) Mapping a V4 Unit’s Receptive Field

4 identical stimuli appear

(200ms)

1 targets cued by luminance increment

(100ms)

Stimuli move to pause position

(750 ms)

Stimuli pause(1000ms)

Stimuli move tofinal position

(750 ms)Cue off(100ms)

Fixation point off: saccade to

the target

Tracking Away from RF

Flashing Gabors+

RF

RFRF

RF

RFRF

RFRF

RFRF

RF

RF

RFRF

4 identical stimuli appear

(200ms)

1 targets cued by luminance increment

(100ms)

Stimuli move to pause position

(750 ms)

Stimuli pause(1000ms)

Stimuli move tofinal position

(750 ms)Cue off(100ms)

Fixation point off: saccade to

the target

Tracking Away from RF

Flashing Gabors+

RFRF

RFRFRFRF

RFRF

RFRFRFRF

RFRFRFRF

RFRFRFRF

RFRF

RFRF

RFRFRFRF

0

100

0

100

0

100

0

100

0

100

0

100

0

100

24681012

24681012

24681012

24681012

24681012

24681012

0

100

Response to Flashed Gabor Image In x,y Space broken down by Orientation

# of Repeats per x,y position

0-33 ms

33-67 ms

67-100 ms

100-133 ms

133-167 ms

167-200 ms

ResponseWindow

TIME

x

y+

x

y+

+1.5

-8.75

-6.25 +11.25

Fixation

x

y+

+1.5

-8.75

-6.25 +11.25

Fixation

Example Neuron:

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24681012

24681012

24681012

24681012

24681012

24681012

0

100

200

# of Repeats per x,y position

0-33 ms

33-67 ms

67-100 ms

100-133 ms

133-167 ms

167-200 ms

TIME

y

x

Response to Flashed Gabor Image In x,y Space broken down by Color

+

Joint Color/Orientation Tuning for Example Cell:

2

4

6

8

2

4

6

8

2

4

6

8

2

4

6

8

2

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8

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0

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1000

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0-33 ms

33-67 ms

67-100 ms

100-133 ms

133-167 ms

167-200 ms

Response Window

TIME

SpatialResponse

Orientation& Color

Response

y

x

# of Repeats

Orientation

Color

X – stat. sig.> baseline

O – stat. sig.< baseline

4 identical stimuli appear

(200ms)

1 or 2 targets cued by luminance increment

(100ms)

Stimuli move to pause position

(750 ms)

Stimuli Pause(1000 ms)

Stimuli move tofinal position

(750 ms)Cue off(100ms)

Fixation point off: saccadeto ALL targets

RFRF

RFRF

RFRF

RFRFRFRF

RFRFRF

RFRF

20

40

60

80

100

Pe

rce

nt

Co

rre

ct

Performance overRecording Sessions

1 of 4Tracking

2 of 4Tracking

Step 2) Preferred Stimuli Enter and Pause in the Unit’s Receptive Field During Tracking

We recorded 29 cells from Monkey M:

1000 1500 2000 2500 3000

10

20

30

40

50

60

70

Time (ms)

Norm

aliz

ed

Response

Mean Response (N=29, All Cells)

Pause Resume

Attended

Unattended

Attended

Unattended

1000 1500 2000 2500 3000

10

20

30

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60

70

Time (ms)

Norm

aliz

ed

Response

Pause Resume

Attended

Unattended

Attended

Unattended

0

10

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40

50

Peak

Norm

aliz

ed

Resp

onse

Modulation = 21.4%

0

10

20

30

40

50

Peak

Norm

aliz

ed

Resp

onse

Modulation = 29.0%

1of4

Tra

ckin

g2

of4

Tra

ckin

g

AI Index1 of 4 Tracking

AI

Ind

ex

2o

f4

Tra

cki

ng

Relative Attentional Modulation for 1 or 2 Target Tracking

-0.2 -0.1 0 0.1 0.2 0.3-0.2

-0.1

0

0.1

0.2

0.3

AI =Attended - Distracter

Attended + Distracter

Corr. = 0.76

NeuronShown Above

* Stat. Sig.

o Non-Sig.

* Stat. Sig.

o Non-Sig.

There were no significant differencesin the strength of modulation for thetracking of a single or two stimuli.

Pause Period

Stimuli pause(75 ms)

Stimuli pause(75 ms)Form Change

(150 ms)

Example Cell:

Unit C20.1: 1 of 4 Tracking

500 1000 1500 2000 2500 3000

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140

500 1000 1500 2000 2500 3000

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140Appear Move Pause Resume End

Tri

alN

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be

r

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Fir

ing

Ra

te(H

z)

Time (ms)

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Fir

ing

Ra

te(H

z)

Target Distracter0

5

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g(

500 1000 1500 2000 2500 3000

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500 1000 1500 2000 2500 3000

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0 500 1000 1500 2000 2500 3000 35000

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be

rF

irin

gR

ate

(Hz)

Time (ms)

Fir

inR

ate

Hz)

Target Distracter

Unit C20.1: 2 of 4 Tracking

Gabor duration 33 msec., One gabor appeared every 8 msec., Resulting in 4 gabors being present at any point in time.

ResponseWindow

Target Distracter

Target Distracter