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The allocation of attention and working memory in visual crowding_

The allocation of attention and working memory in visual crowding_
The allocation of attention and working memory in visual crowding_

The Allocation of Attention and Working Memory

in Visual Crowding

Felix Bacigalupo1,2and Steven J.Luck1

Abstract

■When the distance between a visual target and nearby flankers falls below a critical distance,target discrimination de-clines precipitously.This is called“crowding.”Many researchers have proposed that selective attention plays a role in crowding. However,although some research has examined the effects of directing attention toward versus away from the targets,no pre-vious research has assessed how attentional allocation varies as a function of target–flanker distance in crowding.Here,we used ERPs to assess the operation of attention during crowding, focusing on the attention-related N2pc component.We used a typical crowding task in which participants were asked to report the category(vowel/consonant)of a lateralized target letter flanked by distractor letters at different distances.We tested the hypothesis that attention fails when the target–flanker distance becomes too small for attention to operate effectively.Consistent with this hypothesis,we found that N2pc amplitude was maximal at intermediate target–flanker distances and decreased sub-stantially when crowding became severe.In addition,we exam-ined the sustained posterior contralateral negativity(SPCN), which reflects the amount of information being maintained in working memory.Unlike the N2pc component,the SPCN in-creased in amplitude at small target–flanker distances,suggesting that observers stored information about the target and flankers in working memory when attention failed to select the target. Together,the N2pc and SPCN results suggest that attention and working memory play distinctive roles in crowding:Attention operates to minimize interference from the flankers at inter-mediate target–flanker distances,whereas working memory may be recruited when attention fails to select the target at small target–flanker distances.■

INTRODUCTION

The perception of a peripheral visual target is drama-tically impaired if the target is surrounded by nearby flankers.This phenomenon is called“crowding”(Pelli& Tillman,2008),and it presents a major limit on real-world perception.The mechanisms underlying crowding are hotly debated,and recent reviews have highlighted the potential role of failures of spatial attention(Whitney& Levi,2011;Levi,2008).At intermediate target–distractor distances at the edge of the crowding range,attention may minimize crowding by effectively shrinking neural receptive fields around the target,thereby filtering the flankers(Moran&Desimone,1985).Consistent with this proposal,Yeshurun and Rashal(2010)showed that pre-cuing the target location decreased the target–flanker dis-tance at which crowding occurred.This shows the effect of attention on crowding,but it does not provide any information about how the allocation of attention varies as a function of target–flanker distance(Yeshurun& Rashal,2010).

This study addressed this by recording the N2pc ERP component,which reflects the focusing of attention onto a lateralized target(Hickey,Di Lollo,&McDonald,2009; Luck&Hillyard,1994).N2pc is generated at multiple stages along the ventral processing pathway depending on the scale of target–distractor competition(Hopf et al., 2006),and it is widely used to assess the allocation of attention(Luck,2012;Xu&Franconeri,2012;Hilimire, Mounts,Parks,&Corballis,2009,2010;Eimer&Kiss, 2008;Luck,Girelli,McDermott,&Ford,1997).

We used a typical crowding task in which participants were asked to report the category(vowel/consonant)of a lateralized target letter flanked by distractor letters at dif-ferent distances.Distant distractors were also present in the opposite visual field to create a well-controlled para-digm for measuring the N2pc component.We tested the specific hypothesis that attention can effectively suppress the flankers as long as they are not too close to the target, leading to excellent target discrimination performance. However,when the target–distractor distance becomes too small,the resolution of attention may be insufficient to select the target from the flankers(Intriligator& Cavanagh,2001;He,Cavanagh,&Intriligator,1996),lead-ing to impaired behavioral performance(crowding).If spatial attention is unable to operate effectively at small target–distractor distances,then we should observe both reduced N2pc amplitude and impaired behavioral accu-racy at these distances.

1University of California,Davis,2Pontificia Universidad Catolica

de Chile

?Massachusetts Institute of Technology Journal of Cognitive Neuroscience X:Y,pp.1–14

doi:10.1162/jocn_a_00771

In addition,if spatial attention is unable to select the target from the distractors at small target –flanker dis-tances,then it may be necessary to store information from all of the items in working memory so that the observer can spend the time needed to make an optimal target decision on the basis of the jumble of target and distractor information.This leads to the prediction that more information will be represented in working memory at small target –flanker distances in the period between stimulus offset and the target decision.This should lead to greater amplitude of the sustained posterior contra-lateral negativity (SPCN),which has been shown to in-crease as more items are stored in working memory (Robitaille et al.,2010;Jolicoeur,Brisson,&Robitaille,2008;Vogel &Machizawa,2004).The SPCN has also been reported in visuospatial attention tasks that do not explic-itly require storage of

information in working memory but might still benefit from the maintenance of informa-tion after the stimulus has disappeared (Corriveau et al.,2012;Brisson &Jolicoeur,2007a,2007b).Thus,an in-crease in SPCN amplitude under conditions of crowding would suggest that more items are being maintained in working memory when the distractors cannot be effec-tively filtered.

EXPERIMENT 1

Methods Participants

Fourteen observers (eight women;mean age =20.6years,age range =18–29years)with normal or corrected-to-normal vision and no history of neurological disorders participated.All volunteers gave informed consent and were paid for their participation.The study was approved by the University of California-Davis Institutional Review Board.Two participants were excluded from the analysis,one because of poor performance (correct response rate at chance level in all conditions),reflecting noncompliance with the task instructions,and the other one because of electroencephalographic artifacts on >25%of trials (we always exclude participants for whom >25%of trials are rejected).

Stimuli and Procedure

The stimuli were presented at a distance of 70cm on a cathode ray tube monitor (60-Hz refresh rate)with a black background and a continuously visible white fixa-tion cross.Each stimulus consisted of two vertical arrays of three letters (uppercase Geneva font,height =0.75°),with one array on each side of the fixation cross (see Figure 1).One array was red,and the other was green,varying randomly across trials.The letters on each trial were selected at random without replacement from a set of five vowels (A,E,I,O,and U)and five consonants (N,F,L,G,and J).

Each array was centered 8.8°horizontally from the vertical meridian.The central letter in each array was on the horizontal meridian,with flanker letters directly above and below.The flankers were centered 1.48°,3.13°,or 16.8°from the horizontal meridian.This provided one distance with substantial crowding (1.48°),one dis-tance at the edge of the crowding range (3.13°),and one distance well beyond the crowding range that served as a no-crowding control condition (16.8°).Each distance was equiprobable.Stimulus duration was 200msec,and the SOA varied randomly between 1600and 1800msec.At the beginning of each trial block,the observer was instructed to attend either to red or to green and to indicate whether the middle letter in the array of that color (the target)was a vowel or a consonant.Observers responded by pressing a left or right button depending on whether the target was a vowel or a consonant.Speed and accuracy were equally stressed.Note that this is a conventional design for measuring crowding,except that observers must shift attention to the side containing the attended color to perform the task.The bilateral dis-plays were necessary for measuring N2pc and SPCN with-out contamination from lateralized sensory activity.

Each participant performed 12blocks of 200trials.There were six attend-red and six attend-green blocks,presented in alternating order.Half of the trials were from an additional condition in which the possible flanker locations were arranged in a circular manner rather than in the linear manner shown in Figure 1.However,there

Figure 1.Example stimulus display.Participants fixated on the

central cross while attending to either the red or green items.The side containing the attended color varied unpredictably across trials.The task was to report whether the letter located in the middle of the attended-color array (the target)was a vowel or a consonant.

Experiment 1included three target –flanker distances:16.8°,3.13°,and 1.48°.Experiment 2included no-distractor trials and five target –flanker distances:16.8°,4.12°,3.13°,2.14°,and 1.48°.Conditions included in both experiments are indicated with an asterisk (*)symbol.

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Journal of Cognitive Neuroscience

Volume X,Number Y

was a programming error for this condition,so the data are not reported for these trials.

Recording and Analysis

The EEG was recorded using active Ag–AgCl electrodes (BioSemi ActiveTwo)from the left and right mastoids and32scalp sites(Fp1,Fp2,F7,F3,Fz,F4,F8,T7,C3, Cz,C4,T8,P9,P7,P5,P3,P1,Pz,P2,P4,P6,P8,P10, PO7,PO3,POz,PO4,PO8,O1,Oz,O2,and Iz),according to the modified International10–20System(Picton et al., 2000;American Electroencephalographic Society,1994). To detect eye movements and blinks,the EOG was re-corded from electrodes placed adjacent to the outer canthi of each eye and above and below the right eye. All signals were recorded in single-ended mode and ref-erenced offline.Electrode offsets were kept below 40mV.The EEG and EOG were low-pass filtered with a fifth-order sinc filter(half-power cutoff at208Hz)and digitized at1024Hz.All analyses after data acquisition were conducted using EEGLAB Toolbox(Delorme& Makeig,2004)and ERPLAB Toolbox(Lopez-Calderon& Luck,2014),which are open-source Matlab packages for EEG/ERP analysis.

The EEG signals were referenced offline to the average of the left and right mastoids,and the four EOG signals were referenced offline into bipolar vertical and hori-zontal EOG derivations.These signals were resampled to256Hz(with an antialiasing filter)and filtered offline using a noncausal Butterworth high-pass filter(half-amplitude cutoff=0.1Hz,slope=12dB/octave).A low-pass filter(half-amplitude cutoff=30Hz,slope= 12dB/octave)was used after averaging for plotting pur-poses.Averaged ERP waveforms were computed with an 800-msec epoch,beginning200msec before stimulus onset.

Trials were automatically excluded if the behavioral response was incorrect,if the peak-to-peak voltage ex-ceeded200μV in any200-msec window in any channel, or if a sudden voltage step was detected in the horizontal or vertical EOG(Luck,2005).To assess residual eye movements,we computed averaged horizontal EOG waveforms on left-and right-target trials for each partici-pant,as in previous research(Woodman&Luck,2003). The average residual eye movements in these waveforms were less than3.2μV for all conditions in both experi-ments,corresponding to an eye rotation of less than ±0.1°,with an estimated voltage propagation of less than 0.1μV at the posterior scalp sites(Lins,Picton,Berg,& Scherg,1993).Fewer than25%of trials were rejected owing to artifacts in any of the included participants(mean= 11.5%,range=2.2–22.2%).

The N2pc and SPCN components were measured from contralateral-minus-ipsilateral difference waves(relative to the side of the to-be-identified target)within an a priori ROI consisting of four pairs of electrodes:P5/P6, P7/P8,PO3/PO4,and PO7/PO8.We measured the mean amplitude from these difference waves using a priori time windows of200–300msec for N2pc and400–600msec for SPCN,relative to the200-msec prestimulus baseline period.The waveforms were collapsed across stimulus colors and locations before amplitudes were measured to eliminate sensory confounds related to these factors.

Results and Discussion

Behavior

As shown in Figure2A and Table1,accuracy was high when the target–flanker distance was large but dropped steeply for the most crowded condition.A one-way ANOVA showed a significant effect of distance(F(2,22)= 211.81,p<.001),and follow-up Tukey HSD tests showed significant differences between all pairs of distances(p< .05).RTs exhibited a complementary pattern,increasing as the target–flanker distance decreased(Figure2B and Table1).A one-way ANOVA yielded significant effects of distance(F(2,22)=9.73,p<.001),and Tukey HSD paired tests yielded significant pairwise differences between1.48°and each of the other two distances(3.13°and16.8°). These behavioral results are typical of crowding experi-ments,indicating that our stimuli and task produced normal crowding.

ERPs

N2pc.Figure3shows the ERP waveforms at contralateral and ipsilateral electrode sites,relative to the location of the target,at each target–flanker distance,and Figure4 shows the contralateral-minus-ipsilateral difference waves. In addition,mean amplitude in the N2pc measurement window is shown in Figure2C and Table1.The N2pc can be seen most clearly in the difference waves,where it appears as a negative deflection peaking at approxi-mately225msec.To avoid biasing the statistical analyses by choosing a measurement window on the basis of the observed waveforms,N2pc amplitude was quantified using an a priori time window of200–300msec,which is the canonical period of this component.However, the N2pc was somewhat earlier than that in the present experiment.Thus,to demonstrate that the results of our analyses were not an artifact of the a priori time window of200–300msec,we conducted follow-up analy-ses using windows of150–350,150–320,150–300,and 200–350msec.All of these analyses yielded the same pattern of statistical significance as the a priori window (which was used for all analyses reported below).

As shown in Figures2–4,N2pc amplitude was non-monotonically related to target–flanker distance,with the greatest(most negative)amplitude at a distance of3.13°. Note that this was the same distance at which behavioral accuracy began to deviate from ceiling.A one-way ANOVA

Bacigalupo and Luck3

yielded a significant main effect of target–flanker distance (F(2,22)=4.72,p=.019).The highly nonlinear pattern led to a significant quadratic trend(F(1,11)=4.98,p< .05).This pattern was also confirmed by a Tukey HSD test,which showed

that the N2pc was significantly larger(p<

.05)for the3.13°distance than for the1.48°distance.

These results indicate that maximal attention was allo-

cated at the edge of the crowding range(3.13°),with sig-

nificantly less attention for the most crowded displays

(1.48°).Thus,severe crowding leads to both impaired

attentional allocation and impaired behavioral perfor-

mance.A more detailed discussion of the implications

of the reduced N2pc amplitude will be provided in the

General Discussion.

SPCN.The SPCN can be seen in Figures3and4as a

negativity that begins at approximately350msec and

continues through the end of the epoch.SPCN ampli-

tude was quantified in an a priori time window of400–

600msec,but the same pattern of statistical significance

was obtained with a window of350–600msec.Mean

SPCN amplitude is also shown in Figure2D and Table1.

As shown in Figures2–4,SPCN amplitude increased

monotonically as the target–flanker distance decreased.

A one-way ANOVA yielded a significant main effect of

target–flanker distance(F(2,22)=11.56,p<.001),with

a significant linear trend(F(1,11)=35.73,p<.001).

Post hoc Tukey HSD tests showed significantly larger

SPCN for1.48°compared with both3.13°and16.8°.This

result suggests that working memory,reflected by the

SPCN,is increasingly recruited as crowding increases.As

will be described in more detail in the General Discussion,

it is possible that the SPCN reflects the sustained opera-

tion of attention rather than directly reflects working

memory storage;however,given the brief stimulus dura-

tion,this kind of sustained attention would require that

the information being attended had been stored in work-

ing memory.

Analysis of error trials.The prior analyses were re-

stricted to trials with correct behavioral responses.How-

ever,we also examined the error trials to determine,for

example,whether errors reflected reduced allocation of

attention.The contralateral-minus-ipsilateral difference

waveforms for correct trials and error trials are overlaid

in Figure5.The waveforms were quite noisy on error trials,

especially for the3.13°and16.8°distances,for which

there were few errors(see Table1).There was some

evidence of a reduced N2pc on error trials,especially at

the3.13°distance,but there were no obvious differences

in the SPCN between correct and error trials.To analyze

the data statistically,we conducted two-way ANOVAs on

N2pc and SPCN amplitude with factors of target–distractor

distance and response type(correct vs.error).Neither

the main effect of response type nor the interaction

was significant for either N2pc(F(1,11)=1.75,p=.21;

F(2,22)=2.06,p=.15,respectively)or SPCN(F(1,11)=

0.49,p=.49;F(2,22)=0.05,p=.95,respectively).

We also conducted a comparison of correct and error

trials just at the1.48°distance,where the number of Figure2.Summary of the effect of target–flanker distance on the

behavioral and electrophysiological measures in Experiment1.

(A)Correct response rate.(B)RT.(C)N2pc mean amplitude.

(D)SPCN mean amplitude.Error bars show the within-subject

standard error(Morey,2008).

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error trials was reasonably high.The N2pc was very small on correct trials at this distance,so not surprisingly,there was no significant difference in N2pc amplitude between correct trials and error trials (t (11)=?0.67,p =.51).There was also no significant difference in SPCN ampli-tude

between correct trials and error trials (t (11)=?0.53,p =.61).

Thus,although there was some evidence that the N2pc was reduced on error trials,there was no hint of a reduc-tion in SPCN amplitude on error trials.The SPCN result suggests that errors were not a result of reduced effort

but instead reflect an effect of crowding on the informa-tion available to the neurocognitive process reflected by the SPCN.

EXPERIMENT 2

The goal of Experiment 2was to demonstrate the repli-cability of the results of Experiment 1and to provide a finer manipulation of target –flanker distance.A control condition with no flankers was also included.

Table 1.Mean Accuracy,RT,N2pc Amplitude,SPCN Amplitude,and Number of Error Trials for Each Condition in Experiment 1(SEM in Parentheses)Condition %Correct RT (msec)N2pc (μV)SPCN (μV)Number of Error Trials

16.8°89.6(0.36)654(6.78)?0.32(0.07)?0.58(0.06)30.4(5)3.13°83.8(0.55)682.2(5.84)?0.58(0.06)?0.96(0.07)52(8)1.48°

60(0.48)

721.7(7.84)

?0.08(0.06)

?1.47(0.08)

129.3(9.1)

Figure 3.Grand-averaged contralateral and ipsilateral waveforms relative to the side containing the attended color for Experiment 1.Waveforms from the different target –distractor distances are shown in A (16.8°),B (3.13°),and C (1.48°).The N2pc (200–300msec)and SPCN (400–600msec)measurement windows are highlighted.The waveforms in this and all subsequent ERP figures were low-pass filtered for visual clarity (half-amplitude cutoff =30Hz,slope =12dB/octave).

Bacigalupo and Luck 5

Methods

The methods were identical to those of Experiment 1except as follows.Twenty volunteers (13women;mean age =22.6years,age range =19–32years)with normal or corrected-to-normal vision and no history of neuro-logical disorders participated in this experiment.Two participants were excluded from the analysis because of artifacts on >25%of trials,and two were excluded be-cause of aberrant behavioral performance (unusually low accuracy and fast RTs,indicating a lack of compliance with the task instructions).Fewer than 25%of trials were

Figure 4.Grand-averaged difference waveforms (contralateral minus ipsilateral relative to the side containing the attended color)for the

three target –distractor distances in Experiment 1.The N2pc (200–300msec)and SPCN (400–600msec)measurement windows are highlighted.

Figure 5.Grand-averaged difference waveforms (contralateral minus ipsilateral relative to the side containing the attended color)for correct and error trials in Experiment 1.The N2pc (200–300msec)and SPCN (400–600msec)measurement windows are highlighted.Waveforms from the different target –distractor distances are shown in A (16.8°),B (3.13°),and C (1.48°).

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rejected because of artifacts

in the remaining participants (mean =11.9%,range =2.8–24.7%).

The stimuli and task were identical to those in Exper-iment 1except that flankers were either absent (no distractors)or centered 1.48°,2.14°,3.13°,4.12°,and 16.8°from the horizontal meridian.All stimuli were pre-sented in vertical columns.Results and Discussion Behavior

As shown in Figure 6A and Table 2,accuracy was high when the target –flanker distance was >4°and then fell monotonically at smaller distances.A one-way ANOVA again showed a significant effect of distance (F (5,75)=176.06,p <.001),and follow-up Tukey HSD paired tests showed significant differences between the two most crowded conditions (1.48°vs.2.14°)and also between 1.48°and 2.14°compared with each of the other dis-tances (p <.05).RTs showed the complementary pat-tern,increasing as the target –flanker distance decreased (Figure 6B).A one-way ANOVA yielded a significant effect of distance (F (5,75)=70.91,p <.001).Post hoc Tukey HSD paired tests yielded significant differences between the two most crowded conditions (1.48°vs.2.14°)and also between 1.48°and 2.14°compared with each of the other distances (p <.05).These results replicate the behavioral pattern obtained in Experiment 1and demonstrate that this paradigm yields typical crowding results.ERPs

N2pc.Figure 7shows the ERP waveforms at contralateral and ipsilateral electrode sites,relative to the location of the target,at each target –flanker distance,and Figure 8shows the contralateral-minus-ipsilateral difference waves.For comparison,the difference wave for each target –flanker distance is overlaid with the difference wave for the no-distractor trials.The N2pc appears in the difference waves as a negative deflection peaking between 225and 275msec.As in Experiment 1,N2pc amplitude was quantified using an a priori time window of 200–300msec (see mean am-plitudes in Figure 6C and Table 2),but the same pattern of significance was obtained with windows of 150–350,150–320,150–300,and 200–350msec.

As shown in Figures 6–8and Table 2,the greatest (most negative)N2pc amplitude was found at a target –flanker distance of 3.13°,the same distance as in Experi-ment 1and the distance at which behavioral performance began to be impaired.A one-way ANOVA yielded a sig-nificant main effect of target –flanker distance (F (5,75)=5.1,p <.001).The highly nonlinear pattern led to a sig-nificant quadratic trend (F (1,15)=24.55,p <.001).This quadratic trend was also significant when we excluded the no-distractor condition from the analysis (F (1,15)=9.22,p <.01).This pattern was also confirmed by Tukey

HSD paired tests,which showed that the N2pc was sig-nificantly larger for 3.13°compared with both the most and least crowded conditions (1.48°and no distractors;p <.05).These results replicate the pattern observed in

Figure 6.Summary of the effect of target –flanker distance on the behavioral and electrophysiological measures in Experiment 2.(A)Correct response rate.(B)RT.(C)N2pc mean amplitude.(D)SPCN mean amplitude.Error bars show the within-subject standard error (Morey,2008).

Bacigalupo and Luck

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Experiment1,in which N2pc amplitude was greatest just at the point where behavioral crowding began to occur (3.13°)and declined significantly when the target–distractor distance decreased.This experiment therefore confirms that both attention and behavioral performance are im-paired when the target–flanker distance is sufficiently small. SPCN.The SPCN can be seen in Figures6and7as a negativity that begins at approximately350msec and continues through the end of the epoch.As in Experi-ment1,SPCN amplitude was quantified in an a priori time window of400–600msec,but the same pattern of statistical significance was obtained with a window of 350–600msec.Mean SPCN amplitude is also shown in Figure6D and Table2.

As in Experiment1,we found that SPCN amplitude increased monotonically as the target–flanker distance decreased,yielding a significant main effect of target–flanker distance(F(5,75)=5.81,p<.001)with a signif-icant linear trend(F(1,15)=10.89,p<.01).This linear trend was also significant when we excluded the no-distractor condition from the analysis(F(1,15)=12.54, p<.01).Post hoc Tukey HSD paired tests showed sig-nificantly larger SPCN for the most crowded condition (1.48°)compared with the least crowded ones(4.12°, 16.8°,and no distractors).These results replicate the findings of Experiment1and suggest that working mem-ory is recruited when crowding prevents attention from selecting the target and filtering out the distractors. Analysis of error trials.The contralateral-minus-ipsilateral difference waveforms for correct trials and error trials in Experiment2are overlaid in Figure9.The waveforms were noisier on error trials than on correct trials,espe-cially for the larger target–distractor distances,for which there were few errors(Table2).As in Experiment1, N2pc amplitude was reduced on error trials,especially at distances of3.13°and greater,but there was no consis-tent difference in the SPCN between correct and error trials.To analyze the data statistically,we conducted two-way ANOVAs on N2pc amplitude and SPCN amplitude with factors of target–distractor distance and response type(correct vs.error).For the N2pc,the main effect of response type was significant(F(1,15)=11.71,p=.004), reflecting the reduced N2pc on error trials;the interaction between response type and target–distractor distance was also significant(F(5,75)=2.92,p=.018),reflecting a larger difference between correct and error trials at the larger target–distractor distances.For the SPCN,the main effect of response type was not significant(F(1,15)= 2.47,p=.137),but the interaction was significant(F(5, 75)=2.5,p=.038),reflecting differences between cor-rect and error trials at the intermediate target–distractor distances.

We also conducted a comparison of correct and error trials just at the1.48°distance,where the number of error trials was reasonably high.There was no significant dif-ference between correct trials and error trials for either N2pc amplitude(t(15)=?1.92,p=.07)or SPCN ampli-tude(t(15)=?1.8,p=.09)at this distance.

Although these error-related findings must be consid-ered somewhat speculative given the small number of error trials,the results suggest that errors at large target–distractor distances are accompanied by a failure of the attention mechanism indexed by the N2pc component. This could,in turn,reflect occasional mind wandering or other kinds of off-task performance.It is difficult to draw conclusions about the presence or absence of differences in N2pc between correct and error trials at the small target–distractor distances,however,because the N2pc was small on both correct and error trials at these distances.How-ever,these were the distances at which the SPCN was larg-est,and there was no evidence of a reduced SPCN on error trials at these distances.Thus,when crowding leads to behavioral errors,this does not reflect a reduction in the neurocognitive mechanism reflected by the SPCN but may instead reflect a jumbling of the perceptual infor-mation that is passed to this mechanism by earlier stages of processing.

GENERAL DISCUSSION

Both experiments confirmed the prediction that visuo-spatial attention(as indexed by the N2pc component) would be maximally recruited at intermediate target–flanker distances,near the edge of the crowding range.

Table2.Mean Accuracy,RT,N2pc Amplitude,SPCN Amplitude,and Number of Error Trials for Each Condition in Experiment2 (SEM in Parentheses)

Condition%Correct RT(msec)N2pc(μV)SPCN(μV)Number of Error Trials No distractors93.5(0.19)589.9(3.14)?0.37(0.04)?1.29(0.06)22.8(2.7)

16.8°93.2(0.20)596.6(3.27)?0.65(0.03)?1.20(0.06)23.8(2.7)

4.12°92(0.21)613.6(3.35)?0.73(0.04)?1.17(0.05)27.6(2.9)

3.13°89(0.27)628.2(3.67)?0.89(0.05)?1.55(0.07)35.3(3.6)

2.14°80.3(0.48)67

3.3(3.46)?0.35(0.06)?1.70(0.05)6

4.7(6.1)

1.48°6

2.7(0.43)731(4.93)?0.26(0.05)?1.91(0.05)122.8(6.2)

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Figure7.Grand-averaged contralateral and ipsilateral waveforms relative to the side containing the attended color for Experiment2.The N2pc (200–300msec)and SPCN(400–600msec)measurement windows are highlighted.Waveforms from the different target–distractor distances

are shown in A(no distractors),B(16.8°),C(4.12°),D(3.13°),E(2.14°),and F(1.48°).

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Figure8.Grand-averaged difference waveforms(contralateral minus ipsilateral relative to the side containing the attended color)for Experiment2. The N2pc(200–300msec)and SPCN(400–600msec)measurement windows are highlighted.Waveforms from the different target–distractor distances are shown in A(16.8°),B(4.12°),C(3.13°),D(2.14°),and E(1.48°).For comparison,the waveforms from the no-flanker trials are overlaid with the waveforms for each target–flanker distance.

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Figure9.Grand-averaged difference waveforms(contralateral minus ipsilateral relative to the side containing the attended color)for correct and error trials in Experiment2.The N2pc(200–300msec)and SPCN(400–600msec)measurement windows are highlighted.Waveforms from the different target–distractor distances are shown in A(no distractors),B(16.8°),C(4.12°),D(3.13°),E(2.14°),and F(1.48°).

Bacigalupo and Luck11

This effect was reflected by the finding of maximal N2pc amplitude at3.13°in both experiments.However,when the target–flanker distance decreased from3.13°to1.48°, dramatic changes were observed.1The correct response rate decreased from nearly90%to60%,and N2pc ampli-tude decreased by over50%.These results are consistent with the proposal that attention has a limited spatial reso-lution(Intriligator&Cavanagh,2001;He et al.,1996); after the limit is reached,attention can no longer select the target from the distractors(as reflected in reduced N2pc amplitude),and behavioral performance falls pre-cipitously.This does not rule out the possibility that other factors also contribute to crowding,but it clearly demonstrates that crowding is accompanied by a failure of attention.

Other studies have examined how attention is modu-lated by target–distractor distance.For example,Hilimire et al.(2009,2010)found a linear reduction in N2pc am-plitude according to the distance between the target and a task-relevant distractor(decoy),both of which were distinctive colors presented within a ring of homoge-neously colored distractors.These results are broadly consistent with our findings.However,Hilimire et al.’s (2009,2010)studies did not look at crowding per se, because they varied the distance between two pop-out colors within a ring of distractors,leaving the overall spacing of the stimuli constant.

Yeshurun and Rashal(2010)showed that directing attention toward the target increases performance and reduces the distance at which crowding impacts behav-ioral performance(the critical distance).This finding dovetails perfectly with the present results,which pro-vide direct evidence that the attentional mechanism in-dexed by N2pc amplitude is maximal in the range of target–flanker distances in which behavioral performance is impacted by attention-directing cues.However,the present results go beyond those of Yeshurun and Rashal (2010)by showing how the allocation of attention varies as a function of target–flanker distance.That is,Yeshurun and Rashal’s(2010)results showed the effects of at-tention on perceptual performance,with no way of determining how attentional allocation varied across target–flanker distances.For example,their results are perfectly consistent with an equivalent allocation of atten-tion to the cued side at all target–flanker distances.In contrast,this study demonstrated substantial N2pc ampli-tude at intermediate target–flanker distances relative to no-flanker trials and showed that the N2pc was sig-nificantly reduced when the target–flanker distance fell below a critical distance.

It is important to consider exactly what can be concluded from the reduction in N2pc amplitude observed under conditions of crowding in the present experiment,espe-cially given that it is always challenging to reason backward from a physiological effect to the underlying neurocogni-tive processes(Kappenman&Luck,2012;Poldrack, 2006).There are two main ways in which the N2pc could plausibly be reduced during crowding.First,the reduced N2pc may indicate that participants attempted to focus attention on the target but that the process reflected by N2pc could not work effectively.For example,most of the available evidence indicates that the N2pc component reflects an attentional process that is applied to a target but is needed mainly when competition is present from distrac-tors(Luck,2012;Hickey et al.,2009).Under conditions of crowding,the resolution of attention may be too coarse to allow the target to be selected from among the distractors, and this may result in reduced N2pc amplitude.An alterna-tive,however,is that the visual system detects the crowd-ing preattentively,and participants do not even attempt to focus attention under conditions of severe crowding (which could be either a conscious strategy or an uncon-scious,learned response).In either of these scenarios, however,attention fails to be allocated to the target under conditions of close crowding.Thus,whether partici-pants attempt to focus attention but fail or whether they do not even attempt to focus attention,the present re-sults indicate that the mechanism of attention indexed by the N2pc component is reduced under conditions of crowding.

Whereas the potential role of attention has been wide-ly discussed in the crowding literature(Whitney&Levi, 2011;Levi,2008),we know of no previous reports sug-gesting a role for working memory in visual crowding. However,the present results suggest that working mem-ory was actively involved,with increasing SPCN ampli-tude as the amount of crowding increased.Sustained contralateral negativities are typically associated with the maintenance of information in working memory,with an increasing amplitude as more items are stored in working memory(Robitaille et al.,2010;Jolicoeur et al., 2008;Vogel&Machizawa,2004).The SPCN has also been shown in tasks that do not explicitly ask participants to maintain information online.For example,Corriveau et al.(2012)and Brisson and Jolicoeur(2007a,2007b) reported SPCN activity using a psychological refractory period paradigm,in which participants were asked to se-lect a spatial target without any memory manipulation,and this was interpreted as reflecting the storage of this infor-mation in working memory.

However,sustained contralateral negativities may also be obtained as a result of the sustained focusing of visuo-spatial attention on stimuli that remain visible for long periods,as in the multiple-object tracking paradigm (Drew,Horowitz,Wolfe,&Vogel,2012).Thus,the SPCN observed in this study might reflect the sustained opera-tion of attention rather than reflect working memory storage per se.However,because the stimulus duration in this study was only200msec and the SPCN lasted for at least600msec,information about the stimuli must have been maintained in memory for attention to have operated on them for this long period.Thus,the SPCN effects observed here must be at least an indirect reflec-tion of the storage of information in memory.

12Journal of Cognitive Neuroscience Volume X,Number Y

The present results therefore suggest that working memory,as indexed by the SPCN,is recruited when attention has reached its spatial resolution limit.In other words,when attention cannot focus on the target and filter the distractors,information from multiple items is stored in working memory.The participant can then spend time trying to make an optimal target decision on the basis of this mixture of target and distractor infor-mation.This is consistent with the fact that RTs increased substantially at the small target–distractor distances.The above-chance target discrimination accuracy observed in the most crowded condition may reflect this working memory-based decision process.However,additional research is needed to determine the precise role that working memory plays in crowding.

The analysis of error trials revealed a substantial SPCN component for the most crowded conditions in both ex-periments,without any substantial differences between trials with correct and incorrect responses.These results suggest that participants had successfully stored infor-mation in working memory on error trials and that the errors occurred because this information was noisy (e.g.,a jumbled combination of features of the target and flankers).

In summary,our results suggest that attention and work-ing memory play distinct roles in accomplishing the goal of identifying a target that is surrounded by nearby dis-tractors.When the target is presented without nearby distractors,the need for selective attention and working memory is reduced(although not eliminated).At inter-mediate target–flanker distances,attention is maximally recruited to suppress the flankers and select the target, avoiding a reduction in accuracy that would occur at these distances in the absence of focused attention(Yeshurun &Rashal,2010).When the target–flanker distance falls below the critical distance,attention can no longer select the target and suppress the flankers.At this point,infor-mation from both the target and flankers may be stored in working memory so that the best possible decision can be made from the jumbled perceptual information. Acknowledgments

This study was made possible by Grant R01MH076226to S.J.L. from the National Institute of Mental Health and by MECESUP, VRAID-PUC,and Becas Chile-Conicyt scholarships to F.B.We gratefully acknowledge the assistance of Javier Lopez-Calderon in task programming.F.B.and S.J.L.designed research,F.B. performed research,F.B.analyzed data,and F.B.and S.J.L. wrote the paper.

Reprint requests should be sent to Felix Bacigalupo,Center for Mind and Brain,University of California Davis,267Cousteau Place,Davis,CA95618,or via e-mail:fbacigalupo@https://www.wendangku.net/doc/2811944598.html,. Note

1.Note that the eccentricity of the distractors increased slightly as the target–distractor distance increased.However,most of the behavioral and ERP effects were largest in the comparison with the smallest target–distractor distances(1.48–3.13°),for which the eccentricities were nearly identical(ranging from 8.95°to9.36°).It is very unlikely that this difference in dis-tractor eccentricity could be responsible for the very large dif-ferences in behavioral performance and ERP activity that we observed.

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尊重的素材

尊重的素材(为人处世) 思路 人与人之间只有互相尊重才能友好相处 要让别人尊重自己,首先自己得尊重自己 尊重能减少人与人之间的摩擦 尊重需要理解和宽容 尊重也应坚持原则 尊重能促进社会成员之间的沟通 尊重别人的劳动成果 尊重能巩固友谊 尊重会使合作更愉快 和谐的社会需要彼此间的尊重 名言 施与人,但不要使对方有受施的感觉。帮助人,但给予对方最高的尊重。这是助人的艺术,也是仁爱的情操。—刘墉 卑己而尊人是不好的,尊己而卑人也是不好的。———徐特立 知道他自己尊严的人,他就完全不能尊重别人的尊严。———席勒 真正伟大的人是不压制人也不受人压制的。———纪伯伦 草木是靠着上天的雨露滋长的,但是它们也敢仰望穹苍。———莎士比亚 尊重别人,才能让人尊敬。———笛卡尔 谁自尊,谁就会得到尊重。———巴尔扎克 人应尊敬他自己,并应自视能配得上最高尚的东西。———黑格尔 对人不尊敬,首先就是对自己的不尊敬。———惠特曼

每当人们不尊重我们时,我们总被深深激怒。然而在内心深处,没有一个人十分尊重自己。———马克·吐温 忍辱偷生的人,绝不会受人尊重。———高乃依 敬人者,人恒敬之。———《孟子》 人必自敬,然后人敬之;人必自侮,然后人侮之。———扬雄 不知自爱反是自害。———郑善夫 仁者必敬人。———《荀子》 君子贵人而贱己,先人而后己。———《礼记》 尊严是人类灵魂中不可糟蹋的东西。———古斯曼 对一个人的尊重要达到他所希望的程度,那是困难的。———沃夫格纳 经典素材 1元和200元 (尊重劳动成果) 香港大富豪李嘉诚在下车时不慎将一元钱掉入车下,随即屈身去拾,旁边一服务生看到了,上前帮他拾起了一元钱。李嘉诚收起一元钱后,给了服务生200元酬金。 这里面其实包含了钱以外的价值观念。李嘉诚虽然巨富,但生活俭朴,从不挥霍浪费。他深知亿万资产,都是一元一元挣来的。钱币在他眼中已抽象为一种劳动,而劳动已成为他最重要的生存方式,他的所有财富,都是靠每天20小时以上的劳动堆积起来的。200元酬金,实际上是对劳动的尊重和报答,是不能用金钱衡量的。 富兰克林借书解怨 (尊重别人赢得朋友)

那一刻我感受到了幸福_初中作文

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爸爸唱的英文生日快乐歌虽然不是很动听,但爸爸对我的那份爱我听得很清楚,妈妈为我做的长寿面,我细细的品尝,吃出了爱的味道。妹妹急忙让我许下三个愿望,嘴里不停的唠叨:我知道你的三个愿望是什么?我问:为什么呀!我们是一家人,心连心呀!她高兴的说。 那一刻我才真正解开幸福的密码,感受到了真正的幸福,以前我无法理解幸福,即使身边有够多的幸福也不懂得欣赏,不懂得珍惜,只想拥有更好更贵的,其实幸福比物质更珍贵。 那一刻的幸福就是爱的升华,许多时候能让我们感悟幸福不是名利,物质。而是在血管里涌动着的,漫过心底的爱。 也许每一个人生的那一刻,就是我们幸运的降临在一个温馨的家庭中,而不是降临在孤独的角落里。 家的感觉就是幸福的感觉,幸福一直都存在于我们的身边!

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