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Attentive tracking of multiple objects modulates neuronalresponses in area V4 of macaque.
879
Conclusions
Multiple-Object Tracking Task:
1) Firing rate is enhanced (~18%) in V4 for tracked stimuli.2) The fano factor is decreased (~7.5%) for tracked stimuli.3) The modulation of rate is similar in tracking 1 or 2 items.4) The signal to noise ratio is better in tracking 1 item though.5) Both spike and LFP power spectra are modulated similarly.6) For tracked items, low frequency(5-25Hz) power decreases and high frequency (35-95Hz) power increases.
Jude F. Mitchell, Kristy A. Sundberg & John H. Reynolds
* 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.).
Are V4 Unit’s Rates Modulated?
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
Monkeys track either 1 or 2 of 4 identical items. Stimuluscolor and orientation matched the preferences of a singleunit isolated in V4. During Tracking, a single item entered and paused inside the receptive field for 1000ms. Monkeys indicated the tracked items by making a saccade to each at the end of the trial.
Does the Signal to Noise Ratio Improve for Tracked Items?
H74
Do power spectra of spikes andLFPs change for tracked items?
Attended Target Entering Receptive Field in 2 of 4 Tracking
20
40
60
80-0.2
0.0
+0.2
+0.4
Pause Resume
Fre
quency
(Hz)
500 1000 1500 2000 2500 3000 3500Time (ms)
20
40
60
80 -0.2
0.0
+0.2
+0.4
Distracter Entering Receptive Field in 2 of 4 TrackingPause Resume
Fre
quency
(Hz)
500 1000 1500 2000 2500 3000 3500Time (ms)
20
40
60
80 -0.1
0.0
0.1
500 1000 1500 2000 2500 3000 3500Time (ms)
Pause ResumeAttended Response - Distracter Response in 2 of 4 Tracking
Fre
quency
(Hz)
20
40
60
80 -0.1
0.0
0.1
Attended Response - Distracter Response in 1 of 4 TrackingPause Resume
Fre
quency
(Hz)
500 1000 1500 2000 2500 3000 3500Time (ms)
-0.1
0.0
0.120
40
60
80
500 1000 1500 2000 2500 3000 3500Time (ms)
Pause Resume
Fre
quency
(Hz)
(2 of 4 Attended – Distracter) – (1 of 4 Attended – Distracter)
Attended Target Entering Receptive Field in 2 of 4 Tracking
20
40
60
80-0.2
0.0
+0.2
+0.4
Pause Resume
Fre
quency
(Hz)
500 1000 1500 2000 2500 3000 3500Time (ms)
Attended Target Entering Receptive Field in 2 of 4 Tracking
20
40
60
80-0.2
0.0
+0.2
+0.4
-0.2
0.0
+0.2
+0.4
Pause Resume
Fre
quency
(Hz)
500 1000 1500 2000 2500 3000 3500Time (ms)
500 1000 1500 2000 2500 3000 3500Time (ms)
20
40
60
80 -0.2
0.0
+0.2
+0.4
Distracter Entering Receptive Field in 2 of 4 TrackingPause Resume
Fre
quency
(Hz)
500 1000 1500 2000 2500 3000 3500Time (ms)
20
40
60
80 -0.2
0.0
+0.2
+0.4
-0.2
0.0
+0.2
+0.4
Distracter Entering Receptive Field in 2 of 4 TrackingPause Resume
Fre
quency
(Hz)
500 1000 1500 2000 2500 3000 3500Time (ms)
500 1000 1500 2000 2500 3000 3500Time (ms)
20
40
60
80 -0.1
0.0
0.1
500 1000 1500 2000 2500 3000 3500Time (ms)
Pause ResumeAttended Response - Distracter Response in 2 of 4 Tracking
Fre
quency
(Hz)
20
40
60
80 -0.1
0.0
0.1
-0.1
0.0
0.1
500 1000 1500 2000 2500 3000 3500Time (ms)
500 1000 1500 2000 2500 3000 3500Time (ms)
Pause ResumeAttended Response - Distracter Response in 2 of 4 Tracking
Fre
quency
(Hz)
20
40
60
80 -0.1
0.0
0.1
Attended Response - Distracter Response in 1 of 4 TrackingPause Resume
Fre
quency
(Hz)
500 1000 1500 2000 2500 3000 3500Time (ms)
20
40
60
80 -0.1
0.0
0.1
-0.1
0.0
0.1
Attended Response - Distracter Response in 1 of 4 TrackingPause Resume
Fre
quency
(Hz)
500 1000 1500 2000 2500 3000 3500Time (ms)
500 1000 1500 2000 2500 3000 3500Time (ms)
-0.1
0.0
0.120
40
60
80
500 1000 1500 2000 2500 3000 3500Time (ms)
Pause Resume
Fre
quency
(Hz)
(2 of 4 Attended – Distracter) – (1 of 4 Attended – Distracter)
-0.1
0.0
0.1
-0.1
0.0
0.120
40
60
80
500 1000 1500 2000 2500 3000 3500Time (ms)
500 1000 1500 2000 2500 3000 3500Time (ms)
Pause Resume
Fre
quency
(Hz)
(2 of 4 Attended – Distracter) – (1 of 4 Attended – Distracter)
What is the time course of changein the power spectrum of LFPs?
YES
The Fano Factor Decreasesfor Tracked Items
Changes in Power are not Different Between 1 or 2 Target Tracking
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
Monkeys tracked either 1 or 2 of 4 identical items. Stimuluscolor and orientation were matched to the preferences of anisolated unit in V4. During tracking, an attended or distracteritem entered and paused inside the receptive field for 1000ms. At the end of each trial, the monkey indicated whichitems were tracked by making a saccade to each.
Power spectra were computed using a sliding window of 200mswith a step of 50ms with five Slepian Tapers (W = 6Hz) using Chronux software. Powerwas then normalized at eachfrequency across trial duration and all tracking conditions. Significant attention differences (black dots) were computed, with a Bonferonni correction.
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):1-14.3. Fries P, Reynolds JH, Rorie AE, and Desimone R (2001) Modulation of oscillatory neuronal synchronization by selective visual attention. Science 291:1560-63.
1 2-10
0
10
20
30
Items Tracked
Fir
ing
Ra
teM
od
ula
tion
1 2-10
0
10
20
30
Items Tracked
Fir
ing
Ra
teM
od
ula
tion
Population Summary:
YES
Firing RateModulation is Similarfor Tracking1 or 2 Items
Population Summary:
0 20 40 60 80 1000.95
1.00
1.05
Frequency (Hz)
No
rma
lize
dP
ow
er
Population Spike Spectra in 1 of 4 Tracking
0 20 40 60 80 100
Frequency (Hz)
Population Spike Spectrain 2 of 4 Tracking
AttendedDistracter
AttendedDistracter
1 SEM 1 SEM
N = 118 N = 118Low High Low High
2 1 2-4
-2
0
2
Items Tracked
Mo
du
latio
no
fP
ow
er
1
LowFreq.
HighFreq.
0 20 40 60 80 100
0.6
0.8
1.0
1.2
Frequency (Hz)
No
rma
lize
dP
ow
er
0 20 40 60 80 100
Frequency (Hz)
Population LFP Spectra in 1 of 4 Tracking
Population LFP Spectrain 2 of 4 Tracking
AttendedDistracter
AttendedDistracter
1 SEM
N = 118Low High
1 SEM
N = 118Low High
-30
-20
-10
0
10
20
30
Mo
du
latio
no
fP
ow
er
2 1 2
Items Tracked1
LowFreq.
HighFreq.
*
0 20 40 60 80 1000.95
1.00
1.05
Frequency (Hz)
No
rma
lize
dP
ow
er
Population Spike Spectra in 1 of 4 Tracking
0 20 40 60 80 100
Frequency (Hz)
Population Spike Spectrain 2 of 4 Tracking
AttendedDistracter
AttendedDistracter
1 SEM 1 SEM
N = 118 N = 118Low High Low High
2 1 2-4
-2
0
2
Items Tracked
Mo
du
latio
no
fP
ow
er
1
LowFreq.
HighFreq.
0 20 40 60 80 1000.95
1.00
1.05
Frequency (Hz)
No
rma
lize
dP
ow
er
Population Spike Spectra in 1 of 4 Tracking
0 20 40 60 80 100
Frequency (Hz)
Population Spike Spectrain 2 of 4 Tracking
AttendedDistracter
AttendedDistracter
1 SEM 1 SEM
N = 118 N = 118Low High Low High
0 20 40 60 80 1000.95
1.00
1.05
Frequency (Hz)
No
rma
lize
dP
ow
er
Population Spike Spectra in 1 of 4 Tracking
0 20 40 60 80 100
Frequency (Hz)
Population Spike Spectrain 2 of 4 Tracking
AttendedDistracter
AttendedDistracter
1 SEM 1 SEM
N = 118 N = 118Low High Low High
2 1 2-4
-2
0
2
Items Tracked
Mo
du
latio
no
fP
ow
er
1
LowFreq.
HighFreq.
2 1 2-4
-2
0
2
Items Tracked
Mo
du
latio
no
fP
ow
er
1
LowFreq.
HighFreq.
0 20 40 60 80 100
0.6
0.8
1.0
1.2
Frequency (Hz)
No
rma
lize
dP
ow
er
0 20 40 60 80 100
Frequency (Hz)
Population LFP Spectra in 1 of 4 Tracking
Population LFP Spectrain 2 of 4 Tracking
AttendedDistracter
AttendedDistracter
1 SEM
N = 118Low High
1 SEM
N = 118Low High
-30
-20
-10
0
10
20
30
Mo
du
latio
no
fP
ow
er
2 1 2
Items Tracked1
LowFreq.
HighFreq.
*
0 20 40 60 80 100
0.6
0.8
1.0
1.2
Frequency (Hz)
No
rma
lize
dP
ow
er
0 20 40 60 80 100
Frequency (Hz)
Population LFP Spectra in 1 of 4 Tracking
Population LFP Spectrain 2 of 4 Tracking
AttendedDistracter
AttendedDistracter
1 SEM
N = 118Low High
1 SEM
N = 118Low High
0 20 40 60 80 100
0.6
0.8
1.0
1.2
Frequency (Hz)
No
rma
lize
dP
ow
er
0 20 40 60 80 100
Frequency (Hz)
Population LFP Spectra in 1 of 4 Tracking
Population LFP Spectrain 2 of 4 Tracking
AttendedDistracter
AttendedDistracter
1 SEM
N = 129Low High
1 SEM
N = 129Low High
-30
-20
-10
0
10
20
30
Mo
du
latio
no
fP
ow
er
2 1 2
Items Tracked1
LowFreq.
HighFreq.
*
-30
-20
-10
0
10
20
30
Mo
du
latio
no
fP
ow
er
2 1 2
Items Tracked1
LowFreq.
HighFreq.
*
YES
BothSpike and LFP Power Spectraare Modulated:High Freq. Power Increases andLow Freq. Power Decreases
Population Summary:
Distracter
Attended
Low Frequency Modulation
Distracter
Attended
High Frequency Modulation
0.5 1 1.50.5
1
1.5
N=129
- 1 of 4 Tracking
- 2 of 4 Tracking
0.5 1 1.50.5
1
1.5
- 1 of 4 Tracking
- 2 of 4 Tracking
N=129
Distracter
Attended
Low Frequency Modulation
Distracter
Attended
High Frequency Modulation
0.5 1 1.50.5
1
1.5
N=129
- 1 of 4 Tracking
- 2 of 4 Tracking
0.5 1 1.50.5
1
1.5
0.5 1 1.50.5
1
1.5
N=129
- 1 of 4 Tracking
- 2 of 4 Tracking
- 1 of 4 Tracking
- 2 of 4 Tracking
0.5 1 1.50.5
1
1.5
- 1 of 4 Tracking
- 2 of 4 Tracking
N=129
0.5 1 1.50.5
1
1.5
- 1 of 4 Tracking
- 2 of 4 Tracking
- 1 of 4 Tracking
- 2 of 4 Tracking
N=129
1000 1500 2000 2500 30000
20
40
60
Time (ms)
Pe
ak
No
rma
lize
dR
es
po
ns
e
1000 1500 2000 2500 30000
20
40
60
Time (ms)
Pe
ak
No
rma
lize
dR
es
po
ns
e
Population Mean in 2 of 4 Tracking
Pause Resume
Population Mean in 1 of 4 Tracking
Pause Resume
AttendedDistracter
AttendedDistracter
N=81* N=81*
1 SEM 1 SEM
1000 1500 2000 2500 30000
20
40
60
Time (ms)
Pe
ak
No
rma
lize
dR
es
po
ns
e
1000 1500 2000 2500 30000
20
40
60
Time (ms)
Pe
ak
No
rma
lize
dR
es
po
ns
e
Population Mean in 2 of 4 Tracking
Pause Resume
Population Mean in 1 of 4 Tracking
Pause Resume
AttendedDistracter
AttendedDistracter
N=81* N=81*
1 SEM 1 SEM
1000 1500 2000 2500 3000
0.8
1
1.2
1.4
1.6
1.8
Time (ms)
1000 1500 2000 2500 3000
0.8
1
1.2
1.4
1.6
1.8
Time (ms)
Sp
ike
Va
ria
nce
/M
ea
n
Population Fano Factor in 2 of 4 Tracking
Pause Resume
Sp
ike
Va
rain
ce/
Me
an
Population Fano Factor in 1 of 4 Tracking
Pause Resume
AttendedDistracter
AttendedDistracter
N=81* N=81*
1 SEM 1 SEM
1000 1500 2000 2500 3000
0.8
1
1.2
1.4
1.6
1.8
Time (ms)
1000 1500 2000 2500 3000
0.8
1
1.2
1.4
1.6
1.8
Time (ms)
Sp
ike
Va
ria
nce
/M
ea
n
Population Fano Factor in 2 of 4 Tracking
Pause Resume
Sp
ike
Va
rain
ce/
Me
an
Population Fano Factor in 1 of 4 Tracking
Pause Resume
AttendedDistracter
AttendedDistracter
N=81* N=81*
1 SEM 1 SEM
0 20 40 600
20
40
60
Population Rate Modulationin 1 of 4 Tracking
Attended
(Pe
ak
No
rma
lize
d)
Distracter(Peak Normalized Firing)
N=118(all units)
Monkey A 18.8%Monkey B 16.8%
0 20 40 600
20
40
60
Population Rate Modulationin 2 of 4 Tracking
Attended
(Pe
ak
No
rma
lize
d)
Distracter(Peak Normalized Firing)
N=118(all units)
Monkey A 21.1%Monkey B 16.1%
-50 0 50
-50
0
50
1 of 4 Percent Modulation
2of4
Perc
entM
odula
tion
Correlation of Rate Modulation
N=118(all units)
Monkey A r = 0.91
Monkey B r = 0.70
0 20 40 600
20
40
60
Population Rate Modulationin 1 of 4 Tracking
Attended
(Pe
ak
No
rma
lize
d)
Distracter(Peak Normalized Firing)
N=118(all units)
Monkey A 18.8%Monkey B 16.8%
0 20 40 600
20
40
60
Population Rate Modulationin 1 of 4 Tracking
Attended
(Pe
ak
No
rma
lize
d)
Distracter(Peak Normalized Firing)
N=118(all units)
Monkey A 18.8%Monkey B 16.8%
0 20 40 600
20
40
60
Population Rate Modulationin 2 of 4 Tracking
Attended
(Pe
ak
No
rma
lize
d)
Distracter(Peak Normalized Firing)
N=118(all units)
Monkey A 21.1%Monkey B 16.1%
0 20 40 600
20
40
60
Population Rate Modulationin 2 of 4 Tracking
Attended
(Pe
ak
No
rma
lize
d)
Distracter(Peak Normalized Firing)
N=118(all units)
Monkey A 21.1%Monkey B 16.1%
-50 0 50
-50
0
50
1 of 4 Percent Modulation
2of4
Perc
entM
odula
tion
Correlation of Rate Modulation
N=118(all units)
Monkey A r = 0.91
Monkey B r = 0.70
1 2
1
2
Population Fano Factorin 1 of 4 Tracking
Attended
(Sp
ike
Va
ria
nce
/M
ea
n)
Distracter(Spike Variance / Mean)
N=118(all units)
Monkey A 7.4%Monkey B 12.7%
1 2
1
2
Population Fano Factorin 1 of 4 Tracking
Attended
(Sp
ike
Va
ria
nce
/M
ea
n)
Distracter(Spike Variance / Mean)
N=118(all units)
Monkey A 7.4%Monkey B 12.7%
1 2
1
2
Attended
(Sp
ike
Va
ria
nce
/M
ea
n)
Distracter(Spike Variance / Mean)
Population Fano Factorin 2 of 4 Tracking
Monkey A 4.7%Monkey B 7.8%
N=118(all units)
1 2
1
2
Attended
(Sp
ike
Va
ria
nce
/M
ea
n)
Distracter(Spike Variance / Mean)
Population Fano Factorin 2 of 4 Tracking
Monkey A 4.7%Monkey B 7.8%
N=118(all units)
-40 0 40 80-40
0
40
80Monkey A
r = 0.25Monkey B
r = 0.08
Correlation of Fano FactorModulation
N=118(all units)
2of4
Perc
entM
odula
tion
1 of 4 Percent Modulation
-40 0 40 80-40
0
40
80Monkey A
r = 0.25Monkey B
r = 0.08
Correlation of Fano FactorModulation
N=118(all units)
2of4
Perc
entM
odula
tion
1 of 4 Percent Modulation
1 2-15
-10
-5
0
5
Items Tracked
Fano
Facto
rM
odula
tion
1 2-15
-10
-5
0
5
Items Tracked
Fano
Facto
rM
odula
tion